Dry season conditions determine wet season water use in the wet-tropical savannas of northern Australia

Daily and seasonal patterns of transpiration were measured in evergreen eucalypt trees growing at a wet (Darwin), intermediate (Katherine) and dry site (Newcastle Waters) along a steep rainfall gradient in a north Australian savanna. Relationships between tree size and tree water use were also determined. Diameter at breast height (DBH) was an excellent predictor of sapwood area in the five eucalypt species sampled along the rainfall gradient. A single relationship existed for all species at all sites. Mean daily water use was also correlated to DBH in both wet and dry seasons. There were no significant differences in the relationship between DBH and tree water use at Darwin or Katherine. Among the sites, tree water use was lowest at Newcastle Waters at all DBHs. The relationship between DBH and tree leaf area was similar between species and locations, but the slope of the relationship was less at the end of the dry season than at the end of the wet season at all locations. There was a strong relationship between sapwood area and leaf area that was similar at all sites along the gradient. Transpiration rates were significantly lower in trees at the driest site than at the other sites, but there were no significant differences in transpiration rates between trees growing at Darwin and Katherine. Transpiration rates did not vary significantly between seasons at any site. At all sites, there was only a 10% decline in water use per tree between the wet and dry seasons. A monthly aridity index (pan evaporation/rainfall) and predawn leaf water potential showed strong seasonal patterns. It is proposed that dry season conditions exert control on tree water use during the wet season, possibly through an effect on xylem structure.     

Comparative water use by dryland trees in Parklands in Senegal

Despite the clear evidence of competition for water between trees and crops, there have been very few studies comparing simultaneous water use by differing tree species in drylands. Comparative water use by dryland trees was measured in Senegal using heat balance gauges at the end of the wet season and in the dry season. Significant differences between tree species were found for maximum rates of water use per unit leaf area. Indigenous species may be better adapted to the dry environment than exotic species but the indigenous species Acacia seyal Del. used more water per unit leaf area than all other species. The exotic species Azadirachta indica Adr. Juss. consistently used less water per unit leaf area than most other species. There were significant differences in amounts of water used per unit leaf area by differing provenances of the same tree species. Water use in the dry season varied by a factor of three between two provenances of Acacia aneura F. Muell ex Benth. indicating potential to select provenances for drylands based on their water use characteristics. Absolute rates of water use as well as differences in sapflow between species were greatest when soils were moist suggesting that comparative sapflow studies will be most informative when carried out during the wet season. Water use rankings of the differing tree species were broadly maintained irrespective of season.This revised version was published online in November 2005 with corrections to the Cover Date.     

Contrasting seasonal leaf habits of canopy trees between tropical dry-deciduous and evergreen forests in Thailand

We compared differences in leaf properties, leaf gas exchange and photochemical properties between drought-deciduous and evergreen trees in tropical dry forests, where soil nutrients differed but rainfall was similar. Three canopy trees (Shorea siamensis Miq., Xylia xylocarpa (Roxb.) W. Theob. and Vitex peduncularis Wall. ex Schauer) in a drought-deciduous forest and a canopy tree (Hopea ferrea Lanessan) in an evergreen forest were selected. Soil nutrient availability is lower in the evergreen forest than in the deciduous forest. Compared with the evergreen tree, the deciduous trees had shorter leaf life spans, lower leaf masses per area, higher leaf mass-based nitrogen (N) contents, higher leaf mass-based photosynthetic rates (mass-based P(n)), higher leaf N-based P(n), higher daily maximum stomatal conductance (g(s)) and wider conduits in wood xylem. Mass-based P(n) decreased from the wet to the dry season for all species. Following onset of the dry season, daily maximum g(s) and sensitivity of g(s) to leaf-to-air vapor pressure deficit remained relatively unchanged in the deciduous trees, whereas both properties decreased in the evergreen tree during the dry season. Photochemical capacity and non-photochemical quenching (NPQ) of photosystem II (PSII) also remained relatively unchanged in the deciduous trees even after the onset of the dry season. In contrast, photochemical capacity decreased and NPQ increased in the evergreen tree during the dry season, indicating that the leaves coped with prolonged drought by down-regulating PSII. Thus, the drought-avoidant deciduous species were characterized by high N allocation for leaf carbon assimilation, high water use and photoinhibition avoidance, whereas the drought-tolerant evergreen was characterized by low N allocation for leaf carbon assimilation, conservative water use and photoinhibition tolerance.     

Water economy of Neotropical savanna trees: six paradigms revisited

Biologists have long been puzzled by the striking morphological and anatomical characteristics of Neotropical savanna trees which have large scleromorphic leaves, allocate more than half of their total biomass to belowground structures and produce new leaves during the peak of the dry season. Based on results of ongoing interdisciplinary projects in the savannas of central Brazil (cerrado), we reassessed the validity of six paradigms to account for the water economy of savanna vegetation. (1) All savanna woody species are similar in their ability to take up water from deep soil layers where its availability is relatively constant throughout the year. (2) There is no substantial competition between grasses and trees for water resources during the dry season because grasses exclusively explore upper soil layers, whereas trees access water in deeper soil layers. (3) Tree species have access to abundant groundwater, their stomatal control is weak and they tend to transpire freely. (4) Savanna trees experience increased water deficits during the dry season despite their access to deep soil water. (5) Stomatal conductance of savanna species is low at night to prevent nocturnal transpiration, particularly during the dry season. (6) Savanna tree species can be classified into functional groups according to leaf phenology. We evaluated each paradigm and found differences in the patterns of water uptake between deciduous and evergreen tree species, as well as among evergreen tree species, that have implications for regulation of tree water balance. The absence of resource interactions between herbaceous and woody plants is refuted by our observation that herbaceous plants use water from deep soil layers that is released by deep-rooted trees into the upper soil layer. We obtained evidence of strong stomatal control of transpiration and show that most species exhibit homeostasis in maximum water deficit, with midday water potentials being almost identical in the wet and dry seasons. Although stomatal control is strong during the day, nocturnal transpiration is high during the dry season. Our comparative studies showed that the grouping of species into functional categories is somewhat arbitrary and that ranking species along continuous functional axes better represents the ecological complexity of adaptations of cerrado woody species to their seasonal environment.     

Water-use strategies in two co-occurring Mediterranean evergreen oaks: surviving the summer drought

In the Mediterranean evergreen oak woodlands of southern Portugal, the main tree species are Quercus ilex ssp. rotundifolia Lam. (holm oak) and Quercus suber L. (cork oak). We studied a savannah-type woodland where these species coexist, with the aim of better understanding the mechanisms of tree adaptation to seasonal drought. In both species, seasonal variations in transpiration and predawn leaf water potential showed a maximum in spring followed by a decline through the rainless summer and a recovery with autumn rainfall. Although the observed decrease in predawn leaf water potential in summer indicates soil water depletion, trees maintained transpiration rates above 0.7 mm day(-1) during the summer drought. By that time, more than 70% of the transpired water was being taken from groundwater sources. The daily fluctuations in soil water content suggest that some root uptake of groundwater was mediated through the upper soil layers by hydraulic lift. During the dry season, Q. ilex maintained higher predawn leaf water potentials, canopy conductances and transpiration rates than Q. suber. The higher water status of Q. ilex was likely associated with their deeper root systems compared with Q. suber. Whole-tree hydraulic conductance and minimum midday leaf water potential were lower in Q. ilex, indicating that Q. ilex was more tolerant to drought than Q. suber. Overall, Q. ilex seemed to have more effective drought avoidance and drought tolerance mechanisms than Q. suber.     

Changes in whole-tree water relations during ontogeny of Pinus flexilis and Pinus ponderosa in a high-elevation meadow

We measured sap flux in Pinus ponderosa Laws. and Pinus flexilis James trees in a high-elevation meadow in northern Arizona that has been invaded by conifers over the last 150 years. Sap flux and environmental data were collected from July 1 to September 1, 2000, and used to estimate leaf specific transpiration rate (El), canopy conductance (Gc) and whole-plant hydraulic conductance (Kh). Leaf area to sapwood area ratio (LA/SA) increased with increasing tree size in P. flexilis, but decreased with increasing tree size in P. ponderosa. Both Gc and Kh decreased with increasing tree size in P. flexilis, and showed no clear trends with tree size in P. ponderosa. For both species, Gc was lower in the summer dry season than in the summer rainy season, but El did not change between wet and dry summer seasons. Midday water potential (Psi(mid)) did not change across seasons for either species, whereas predawn water potential (Psi(pre)) tracked variation in soil water content across seasons. Pinus flexilis showed greater stomatal response to vapor pressure deficit (VPD) and maintained higher Psi(mid) than P. ponderosa. Both species showed greater sensitivity to VPD at high photosynthetically active radiation (PAR; > 2500 micromol m-2 s-1) than at low PAR (< 2500 micromol m-2 s-1). We conclude that the direction of change in Gc and Kh with increasing tree size differed between co-occurring Pinus species, and was influenced by changes in LA/SA. Whole-tree water use and El were similar between wet and dry summer seasons, possibly because of tight stomatal control over water loss.     

Functional convergence in hydraulic architecture and water relations of tropical savanna trees: from leaf to whole plant

Functional convergence in hydraulic architecture and water relations, and potential trade-offs in resource allocation were investigated in six dominant neotropical savanna tree species from central Brazil during the peak of the dry season. Common relationships between wood density and several aspects of plant water relations and hydraulic architecture were observed. All species and individuals shared the same negative exponential relationship between sapwood saturated water content and wood density. Wood density was a good predictor of minimum (midday) leaf water potential and total daily transpiration, both of which decreased linearly with increasing wood density for all individuals and species. With respect to hydraulic architecture, specific and leaf-specific hydraulic conductivity decreased and the leaf:sapwood area ratio increased more than 5-fold as wood density increased from 0.37 to 0.71 g cm(-3) for all individuals and species. Wood density was also a good predictor of the temporal dynamics of water flow in stems, with the time of onset of sap flow in the morning and the maximum sap flow tending to occur progressively earlier in the day as wood density increased. Leaf properties associated with wood density included stomatal conductance, specific leaf area, and osmotic potential at the turgor loss point, which decreased linearly with increasing wood density. Wood density increased linearly with decreasing bulk soil water potential experienced by individual plants during the dry season, suggesting that wood density was greatest in individuals with mostly shallow roots, and therefore limited access to more abundant soil water at greater depths. Despite their taxonomic diversity and large intrapopulation differences in architectural traits, the six co-occurring species and their individuals shared similar functional relationships between all pairs of variables studied. Thus, rather than differing intrinsically in physiological responsiveness, the species and the individuals appeared to have distinct operating ranges along common physiological response curves dictated by plant architectural and structural features. The patterns of water uptake and access to soil water during the dry season appeared to be the main determinant of wood density, which constrained evolutionary options related to plant water economy and hydraulic architecture, leading to functional convergence in the neotropical savanna trees studied.     

Stable annual pattern of water use by Acacia tortilis in Sahelian Africa

Water use by mature trees of Acacia tortilis (Forsk.) Hayne ssp. raddiana (Savi) Brenan var. raddiana growing in the northern Sahel was continuously recorded over 4 years. Water use was estimated from xylem sap flow measured by transient heat dissipation. Concurrently, cambial growth, canopy phenology, leaf water potential, climatic conditions and soil water availability (SWA) were monitored. In addition to the variation attributable to interannual variation in rainfall, SWA was increased by irrigation during one wet season. The wet season lasted from July to September, and annual rainfall ranged between 146 and 367 mm. The annual amount and pattern of tree water use were stable from year-to-year despite interannual and seasonal variations in SWA in the upper soil layers. Acacia tortilis transpired readily throughout the year, except for one month during the dry season when defoliation was at a maximum. Maximum water use of about 23 l (dm sapwood area)(-2) day(-1) was recorded at the end of the wet season. While trees retained foliage in the dry season, the decline in water use was modest at around 30%. Variation in predawn leaf water potential indicated that the trees were subject to soil water constraint. The rapid depletion of water in the uppermost soil layers after the wet season implies that there was extensive use of water from deep soil layers. The deep soil profile revealed (1) the existence of living roots at 25 m and (2) that the availability of soil water was low (-1.6 MPa) down to the water table at a depth of 31 m. However, transpiration was recorded at a predawn leaf water potential of -2.0 MPa, indicating that the trees used water from both intermediary soil layers and the water table. During the full canopy stage, mean values of whole-tree hydraulic conductance were similar in the wet and dry seasons. We propose that the stability of water use at the seasonal and annual scales resulted from a combination of features, including an extensive rooting habit related to deep water availability and an effective regulation of canopy conductance. Despite a limited effect on tree water use, irrigation during the wet season sharply increased predawn leaf water potential and cambial growth of trunks and branches.     

Transpiration increases during the dry season: patterns of tree water use in eucalypt open-forests of northern Australia

Australian savannas exhibit marked seasonality in precipitation, with more than 90% of the annual total falling between October and May. The dry season is characterized by declining soil water availability and high vapor pressure deficits (up to 2.5 kPa). We used heat pulse technology to measure whole-tree transpiration rates on a daily and seasonal basis for the two dominant eucalypts at a site near Darwin, Australia. Contrary to expectations, transpiration rates were higher during the dry season than during the wet season, largely because of increased evaporative demand and the exploitation of groundwater reserves by the trees. Transpiration rates exhibited a marked hysteresis in relation to vapor pressure deficit, which was more marked in the dry season than in the wet season. This result may be attributable to low soil hydraulic conductivity, or the use of stored stem water, or both. Tree water use was strongly correlated with leaf area and diameter at breast height and there were no differences in transpiration between the species studied. These results are discussed in relation to scaling tree water use to stand water use.     

Gender-specific patterns of aboveground allocation, canopy conductance and water use in a dominant riparian tree species: Acer negundo

Acer negundo Sarg. (box elder) is a dioecious tree species that dominates riparian systems at mid elevations throughout the southwest and Intermountain West of the United States. Previous studies have shown that female A. negundo trees occur at higher frequencies along stream margins, whereas males occur at higher frequencies in drier microsites. To better understand the adaptive significance of sex ratio biases and their impact on the ecohydrology of riparian ecosystems, we examined whole-plant water relations and hydraulic properties of mature male and female A. negundo trees occurring within 1 m of a perennial stream channel. We hypothesized that (1) females would have significantly greater canopy water fluxes than males (particularly during periods of seed production: May-June), and (2) xylem in females is more hydraulically efficient but more vulnerable to cavitation than xylem in males. Mean sap flux density (J(s)) during the early growing season (May and June) was 43% higher in female trees than in male trees (n = 6 and 7 trees respectively, P < 0.0001). Mean J(s) in July and August remained 17% higher in females than in males (P = 0.0009). Mean canopy stomatal conductance per unit leaf area (g(s,leaf)) in May and June was on average 140% higher in females than in males (P < 0.0001). Mean g(s,leaf) in July and August remained 69% higher in female trees than in male trees (P < 0.0001). Canopy stomatal conductance scaled to basal area was 90 and 31% higher in females relative to males during May-June and July-August, respectively (P < 0.0001 during both periods). Conversely, there were no apparent differences in either branch hydraulic conductance or branch xylem cavitation vulnerability between genders. These results improve our capacity to describe the adaptive forces that shape the spatial distribution of male and female trees in dioecious species, and their consequences for ecohydrological processes in riparian ecosystems.     

Photosynthesis and water relations of savanna tree species differing in leaf phenology

Godmania macrocarpa Hemsley, a deciduous tree characteristic of fire protected areas of the savanna region of central Venezuela, was more drought tolerant, allocated more N to leaves and had consistently higher photosynthetic rates than Curatella americana L., a ubiquitous species growing successfully within the grasslands of tropical American savannas. Godmania macrocarpa maintained higher leaf conductance and photosynthesized at higher xylem water tensions than C. americana. As the dry season progressed, G. macrocarpa was more affected by water stress than C. americana, which may explain why G. macrocarpa shed its leaves before forming new leaves. For both species, leaf sap osmolality was strongly correlated with, but not completely accounted for by, soluble sugars. Integrated water-use efficiency, as measured by delta(13)C, was similar for both species, but young leaves were more efficient than old leaves. Water-use efficiency of adult leaves was similar in both species as a result of higher photosynthetic rates in G. macrocarpa and lower leaf conductances in C. americana. Compared to G. macrocarpa, instantaneous photosynthetic N-use efficiency was higher in C. americana despite its lower maximum photosynthetic rates. The absence of G. macrocarpa trees from open grasslands, despite their high productive capacity, is possibly the result of unfavorable conditions for germination, poor survival of seedlings, and lack of resistance against fire.     

Transport,storage and mobilization of nitrogen by trees and shrubs in the wet/dry tropics of northern Australia

Xylem sap from woody species in the wet/dry tropics of northern Australia was analyzed for N compounds. At the peak of the dry season, arginine was the main N compound in sap of most species of woodlands and deciduous monsoon forest. In the wet season, a marked change occurred with amides becoming the main sap N constituents of most species. Species from an evergreen monsoon forest, with a permanent water source, transported amides in the dry season. In the dry season, nitrate accounted for 7 and 12% of total xylem sap N in species of deciduous and evergreen monsoon forests, respectively. In the wet season, the proportion of N present as nitrate increased to 22% in deciduous monsoon forest species. These results suggest that N is taken up and assimilated mainly in the wet season and that this newly assimilated N is mostly transported as amide-N (woodland species, monsoon forest species) and nitrate (monsoon forest species). Arginine is the form in which stored N is remobilized and transported by woodland and deciduous monsoon forest species in the dry season. Several proteins, which may represent bark storage proteins, were detected in inner bark tissue from a range of trees in the dry season, indicating that, although N uptake appears to be limited in the dry season, the many tree and shrub species that produce flowers, fruit or leaves in the dry season use stored N to support growth. Nitrogen characteristics of the studied species are discussed in relation to the tropical environment.     

The annual pattern of sap flow in two Eucalyptus species established in the vicinity of gold-mine tailings dams in central South Africa

Several hundred mine tailings dams occur in the Witwatersrand Basin Goldfields in central South Africa. Seepage of acid mine drainage (AMD) from these unlined structures is widespread, and a variety of contaminants is released into soil and groundwater. The ‘Mine Woodlands Project’ is aimed at evaluating the use of high-density tree stands over surrounding contaminant plumes to limit the spread of contaminants through hydraulic control of groundwater and enhanced uptake or immobilisation of contaminants. The annual pattern of hourly sap flow in four contiguous Eucalyptus dunnii trees (aged three years) was followed over a full year in a species trial situated near Carltonville. The annual pattern of hourly sap flow was also recorded in four contiguous sample trees (aged four years) of the clonal hybrid E. grandis × E. camaldulensis (E. G×C) at another trial near Orkney. Both species showed high sap flow rates close to reference evaporation rates in response to summer rains. Both showed greatly reduced sap flow rates during the latter half of the dry winter season. Sap flow rates only recovered after the arrival of the first spring rains. Annual sap flow (E. dunnii, 673 mm; E. G×C, 767 mm) was similar to the recorded annual rainfall at each site (E. dunnii, 629 mm; E. G×C, 795 mm), and was substantially lower than total annual reference evaporation (E. dunnii, 1 273 mm; E. G×C, 1 330 mm). We conclude that the roots of both species are not yet deep enough to access the AMD-influenced groundwater, which lies at depths of 14 and 10 m below the ground at the Carltonville and Orkney sites, respectively. Despite prolonged water deficits, both species survived well and maintained sufficient vigour to permit the quick recovery of high transpiration rates in the following summer. This resilience is essential to hasten root growth and improve the chance of contact with groundwater plumes.     

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.     

Impact of soil drought on sap flow and water status of evergreen trees in a tropical monsoon forest in northern Thailand

Hill evergreen forest is the dominant vegetation type in northern Thailand. In this region, there is higher atmospheric evaporative demand and lower soil moisture during the 5- to 7-month dry season than in the rainy season under influences from Asian monsoons. In an earlier study we revealed that canopy-scale transpiration is actively maintained even during the latter part of the dry season in hill evergreen forest. However, the impact of soil drought on tree water use was not investigated. To clarify the ecohydrological processes at this site, we used individual tree-scale measurements during a 2-year period to base our examination of whether limited water use in individual trees is caused by soil drought in the latter part of the dry season. Sap flow and water potential measurements were conducted in four evergreen trees, two large emergent trees 29.8 and 25.4 m high, and two smaller understory trees 4.8 and 1.4 m high.The amount of rainfall preceding the late dry season of 2004 was significantly less than that preceding the late dry season of 2003. Although a distinct decrease in sap-flow velocities in individual trees due to soil water stress was not found in the late dry season of 2003, it did become comparatively apparent in the late dry season of 2004; ranging from 10 to 40% for a given atmospheric evaporative demand. Furthermore, the reductions in sap-flow velocities and predawn stem-water potential were most significant in the smallest tree. The recovery of sap-flow velocities and water potential in the smallest tree after irrigation confirmed that the reductions in sap-flow velocity and predawn stem-water potential in the smallest tree were caused by soil drought. These results suggest that shallower roots could be reason for the significant decrease in water use in the smallest trees. The deeper roots of larger trees could be the reason for the reduced impact of soil drought on water use in larger trees, and canopy-scale transpiration might be maintained by larger trees, even in an unusually severe drought. These possibilities provide a new insight for management of evergreen forests under Asian monsoon influences.     

Electric resistance as a measure of tree water status during seasonal drought in a tropical dry forest in Costa Rica

Variation in electric resistance of stem tissues was used to measure differences and changes in water status among trees in a tropical dry forest in Costa Rica during the dry season. For more than 30 tree species, stem water content (SWC), measured as electric resistance between nails driven 20 mm deep into tree trunks, correlated well with wood density, saturation water content, dehydration, measured with the pressure chamber, and tree development during drought. At dry sites, SWC was lowest in hardwood trees (characterized by high wood density) and highest in stem-succulent lightwood trees (characterized by low wood density). Among hardwood trees, SWC varied with soil water availability. During the dry season, SWC declined before leaf shedding and increased during rehydration preceding bud break. The time course of seasonal changes in SWC apparently constitutes an indirect measure of variation in the relative water content of outer stem tissues, which determines development of dry-forest trees during the dry season.     

Responses of Acer negundo genders to interannual differences in water availability determined from carbon isotope ratios of tree ring cellulose

Understanding the responses of riparian trees to water availability is critical for predicting the effects of changes in precipitation on riparian ecosystems. Dioecious Acer negundo L. (box elder) is a common riparian tree that is highly sensitive to water stress. Earlier studies indicated that the genders of A. negundo respond differently to water availability, with males being more conservative in their water use than females. To assess the potential effects of changes in precipitation on the sex ratio of riparian trees, we extended earlier studies of A. negundo by analyzing responses of male and female genotypes to interannual differences in water availability in a common garden. We measured growth of tree rings and used stable carbon isotope analysis of tree ring alpha-cellulose to integrate physiological responses to annual water treatments. During dry years, male and female trees exhibited similar growth and physiological responses. However, during wet years, females exhibited higher growth rates and lower carbon isotope ratios (indicating less conservative water use) than did males. Furthermore, we found that male trees exhibited similar stomatal behavior (inferred from integrated carbon isotope ratios) whether years were wet or dry, whereas females did not exhibit a consistent response to changes in water availability. We predict that with increasing precipitation and soil water availability, the representation of females will be favored because of shifts in the competitive interactions of the genders. Such changes may affect the reproductive output of these riparian trees and may influence overall water flux from riparian ecosystems. In addition, this study demonstrates the utility of carbon isotope analysis for assessing long-term responses of tree populations to shifts in water availability.     

Ecophysiology of Acacia species in wet–dry tropical plantations

Selected tropical Acacia species are used extensively for short-rotation plantation forestry in many parts of Asia and, to a limited degree, in Australia. We explored leaf-level photosynthetic activity and leaf water potential (Ψleaf) of three field-grown Acacia tree species (aged between 7 and 18 months) in contrasting wet–dry tropical plantations in southern Vietnam and northern Australia. Light-saturated photosynthetic rate (A1500) declined throughout the morning and early afternoon in the dry season; in the wet season, levels remained high and relatively constant throughout most of the day. Maximum daily A1500 at 09:00 ranged from 22.2 μmol?m?2?s?1 in the wet to 10.4 μmol?m?2?s?1 in the dry season. At both locations, trees were able to extract soil water such that pre-dawn leaf water potential (Ψpd) remained>?1.5?MPa even at the end of the dry season. Stomatal conductance to water vapour (gs) did not respond to decreasing Ψleaf during the wet season but was sensitive to changes in Ψleaf in the dry season. Species comparisons of the relationships between A1500 and Ψleaf revealed different strategies to balance carbon uptake and water loss in a wet–dry environment. Acacia crassicarpa and A. mangium regulated Ψleaf to a greater extent than the A. mangium×A. auriculiformis hybrid such that ?Ψleaf (determined as Ψpd?midday Ψleaf) was unaffected by season. This result suggests that the hydraulic regulation of tree water status varies amongst young tropical Acacia species. From a management perspective, for Acacia species that tend to strongly regulate water loss in environments with an extended dry season, overall productivity at the end of a rotation may be less than for species that prioritise carbon gain.     

Dynamics of transpiration, sap flow and use of stored water in tropical forest canopy trees

In large trees, the daily onset of transpiration causes water to be withdrawn from internal storage compartments, resulting in lags between changes in transpiration and sap flow at the base of the tree. We measured time courses of sap flow, hydraulic resistance, plant water potential and stomatal resistance in co-occurring tropical forest canopy trees with trunk diameters ranging from 0.34-0.98 m, to determine how total daily water use and daily reliance on stored water scaled with size. We also examined the effects of scale and tree hydraulic properties on apparent time constants for changes in transpiration and water flow in response to fluctuating environmental variables. Time constants for water movement were estimated from whole-tree hydraulic resistance (R) and capacitance (C) using an electric circuit analogy, and from rates of change in water movement through intact trees. Total daily water use and reliance on stored water were strongly correlated with trunk diameter, independent of species. Although total daily withdrawal of water from internal storage increased with tree size, its relative contribution to the daily water budget (approximately 10%) remained constant. Net withdrawal of water from storage ceased when upper branch water potential corresponded to the sapwood water potential (Psi(sw)) at which further withdrawal of water from sapwood would have caused Psi(sw) to decline precipitously. Stomatal coordination of vapor and liquid phase resistances played a key role in limiting stored water use to a nearly constant fraction of total daily water use. Time constants for changes in transpiration, estimated as the product of whole- tree R and C, were similar among individuals (~0.53 h), indicating that R and C co-varied with tree size in an inverse manner. Similarly, time constants estimated from rates of change in crown and basal sap flux were nearly identical among individuals and therefore independent of tree size and species.     

Estimating stand water use of large mountain ash trees and validation of the sap flow measurement technique

Mountain ash (Eucalyptus regnans F.J. Muell.) forest catchments exhibit a strong relationship between stand age and runoff, attributed inter alia to differences in tree water use. However, the tree water use component of the mountain ash forest water balance is poorly quantified. We have used the sap flow technique to obtain estimates of daily water use in large mountain ash trees. First, the sap flow technique was validated by means of an in situ cut tree experiment. Close agreement was obtained between the sap flow estimate of water use and the actual uptake of water by the tree from a reservoir. Second, we compared the variability in sap velocity between a symmetric and an asymmetric tree by using multiple sap flow loggers. In the symmetric tree, velocity was fairly uniform throughout the xylem during the day, indicating that accurate sap flow estimates can be obtained with a minimal number of sampling points. However, large variations in sap velocity were observed in the asymmetric tree, indicating that much larger sampling sizes are required in asymmetric stems for an accurate determination of mean sap velocity. Finally, we compared two procedures for scaling individual tree sap flow estimates to the stand level based on stem diameter and leaf area index measurements. The first procedure was based on a regression between stem diameter and tree water use, developed on a small sample of trees and applied to a stand-level census of stem diameter values. Inputs to the second procedure were tree water use and leaf area of a single tree and the leaf area index of the stand. The two procedures yielded similar results; however, the first procedure was more robust but it required more sampling effort than the second procedure.