Water-use efficiency in a semi-arid woodland with high rainfall variability.

As the ratio of carbon uptake to water use by vegetation, water-use efficiency (WUE) is a key ecosystem property linking global carbon and water cycles. It can be estimated in several ways, but it is currently unclear how different measures of WUE relate, and how well they each capture variation in WUE with soil moisture availability. We evaluated WUE in an Acacia-dominated woodland ecosystem of central Australia at various spatial and temporal scales using stable carbon isotope analysis, leaf gas exchange and eddy covariance (EC) fluxes. Semi-arid Australia has a highly variable rainfall pattern, making it an ideal system to study how WUE varies with water availability. We normalized our measures of WUE across a range of vapour pressure deficits using g1 , which is a parameter derived from an optimal stomatal conductance model and which is inversely related to WUE. Continuous measures of whole-ecosystem g1 obtained from EC data were elevated in the 3 days following rain, indicating a strong effect of soil evaporation. Once these values were removed, a close relationship of g1 with soil moisture content was observed. Leaf-scale values of g1 derived from gas exchange were in close agreement with ecosystem-scale values. In contrast, values of g1 obtained from stable isotopes did not vary with soil moisture availability, potentially indicating remobilization of stored carbon during dry periods. Our comprehensive comparison of alternative measures of WUE shows the importance of stomatal control of fluxes in this highly variable rainfall climate and demonstrates the ability of these different measures to quantify this effect. Our study provides the empirical evidence required to better predict the dynamic carbon-water relations in semi-arid Australian ecosystems.

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.

Effectiveness of time of sowing and cultivar choice for managing climate change: wheat crop phenology and water use efficiency.

Climate change (CC) presents a challenge for the sustainable development of wheat production systems in Australia. This study aimed to (1) quantify the impact of future CC on wheat grain yield for the period centred on 2030 from the perspectives of wheat phenology, water use and water use efficiency (WUE) and (2) evaluate the effectiveness of changing sowing times and cultivars in response to the expected impacts of future CC on wheat grain yield. The daily outputs of CSIRO Conformal-Cubic Atmospheric Model for baseline and future periods were used by a stochastic weather generator to derive changes in mean climate and in climate variability and to construct local climate scenarios, which were then coupled with a wheat crop model to achieve the two research aims. We considered three locations in New South Wales, Australia, six times of sowing (TOS) and three bread wheat (Triticum aestivum L.) cultivars in this study. Simulation results show that in 2030 (1) for impact analysis, wheat phenological events are expected to occur earlier and crop water use is expected to decrease across all cases (the combination of three locations, six TOS and three cultivars), wheat grain yield would increase or decrease depending on locations and TOS; and WUE would increase in most of the cases; (2) for adaptation considerations, the combination of TOS and cultivars with the highest yield varied across locations. Wheat growers at different locations will require different strategies in managing the negative impacts or taking the opportunities of future CC.

Long-term Water Table Monitoring of Rio Grande Riparian Ecosystems for Restoration Potential Amid Hydroclimatic Challenges.

Hydrological processes drive the ecological functioning and sustainability of cottonwood-dominated riparian ecosystems in the arid southwestern USA. Snowmelt runoff elevates groundwater levels and inundates floodplains, which promotes cottonwood germination. Once established, these phreatophytes rely on accessible water tables (WTs). In New Mexico's Middle Rio Grande corridor diminished flooding and deepening WTs threaten native riparian communities. We monitored surface flows and riparian WTs for up to 14 years, which revealed that WTs and surface flows, including peak snowmelt discharge, respond to basin climate conditions and resource management. WT hydrographs influence the composition of riparian communities and can be used to assess if potential restoration sites meet native vegetation tolerances for WT depths, rates of recession, and variability throughout their life stages. WTs were highly variable in some sites, which can preclude native vegetation less adapted to deep drawdowns during extended droughts. Rates of WT recession varied between sites and should be assessed in regard to recruitment potential. Locations with relatively shallow WTs and limited variability are likely to be more viable for successful restoration. Suitable sites have diminished greatly as the once meandering Rio Grande has been constrained and depleted. Increasing demands on water and the presence of invasive vegetation better adapted to the altered hydrologic regime further impact native riparian communities. Long-term monitoring over a range of sites and hydroclimatic extremes reveals attributes that can be evaluated for restoration potential.

Responses of LAI to rainfall explain contrasting sensitivities to carbon uptake between forest and non-forest ecosystems in Australia.

Non-forest ecosystems (predominant in semi-arid and arid regions) contribute significantly to the increasing trend and interannual variation of land carbon uptake over the last three decades, yet the mechanisms are poorly understood. By analysing the flux measurements from 23 ecosystems in Australia, we found the the correlation between gross primary production (GPP) and ecosystem respiration (Re) was significant for non-forest ecosystems, but was not for forests. In non-forest ecosystems, both GPP and Re increased with rainfall, and, consequently net ecosystem production (NEP) increased with rainfall. In forest ecosystems, GPP and Re were insensitive to rainfall. Furthermore sensitivity of GPP to rainfall was dominated by the rainfall-driven variation of LAI rather GPP per unit LAI in non-forest ecosystems, which was not correctly reproduced by current land models, indicating that the mechanisms underlying the response of LAI to rainfall should be targeted for future model development.

Quantifying deforestation and forest degradation with thermal response.

Deforestation and forest degradation cause the deterioration of resources and ecosystem services. However, there are still no operational indicators to measure forest status, especially for forest degradation. In the present study, we analysed the thermal response number (TRN, calculated by daily total net radiation divided by daily temperature range) of 163 sites including mature forest, disturbed forest, planted forest, shrubland, grassland, savanna vegetation and cropland. TRN generally increased with latitude, however the regression of TRN against latitude differed among vegetation types. Mature forests are superior as thermal buffers, and had significantly higher TRN than disturbed and planted forests. There was a clear boundary between TRN of forest and non-forest vegetation (i.e. grassland and savanna) with the exception of shrubland, whose TRN overlapped with that of forest vegetation. We propose to use the TRN of local mature forest as the optimal TRN (TRNopt). A forest with lower than 75% of TRNopt was identified as subjected to significant disturbance, and forests with 66% of TRNopt was the threshold for deforestation within the absolute latitude from 30° to 55°. Our results emphasized the irreplaceable thermal buffer capacity of mature forest. TRN can be used for early warning of deforestation and degradation risk. It is therefore a valuable tool in the effort to protect forests and prevent deforestation.

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.

Variation in photosynthetic traits related to access to water in semiarid Australian woody species.

Low soil water content can limit photosynthesis by reducing stomatal conductance. Here, we explore relationships among traits pertaining to carbon uptake and pre-dawn leaf water potential (as an index of soil water availability) across eight species found in semiarid central Australia. We found that as pre-dawn leaf water potential declined, stomatal limitations to photosynthesis increased, as did foliar nitrogen, which enhanced photosynthesis. Nitrogen-fixing Acacia species had higher foliar nitrogen concentrations compared with non-nitrogen fixing species, although there was considerable variability of traits within the Acacia genus. From principal component analysis we found that the most dissimilar species was Acacia aptaneura Maslin&J.E.Reid compared with both Eucalyptus camaldulensis Dehnh. and Corymbia opaca. (D.J.Carr & S.G.M.Carr)K.D.Hill&L.A.S.Johnson, having both the largest foliar N content, equal largest leaf mass per area and experiencing the lowest pre-dawn water potential of all species. A. aptaneura has shallow roots and grows above a hardpan that excludes access to groundwater, in contrast to E. camaldulensis and C. opaca, which are known to access groundwater. We conclude that ecohydrological niche separation is an important factor driving the variability of within-biome traits related to carbon gain. These observations have important implications for global vegetation models, which are parameterised with many of the traits measured here, but are often limited by data availability.

Divergence in plant water-use strategies in semiarid woody species.

Partitioning of water resources amongst plant species within a single climate envelope is possible if the species differ in key hydraulic traits. We examined 11 bivariate trait relationships across nine woody species found in the Ti-Tree basin of central Australia. We found that species with limited access to soil moisture, evidenced by low pre-dawn leaf water potential, displayed anisohydric behaviour (e.g. large seasonal fluctuations in minimum leaf water potential), had greater sapwood density and lower osmotic potential at full turgor. Osmotic potential at full turgor was positively correlated with the leaf water potential at turgor loss, which was, in turn, positively correlated with the water potential at incipient stomatal closure. We also observed divergent behaviour in two species of Mulga, a complex of closely related Acacia species which range from tall shrubs to low trees and dominate large areas of arid and semiarid Australia. These Mulga species had much lower minimum leaf water potentials and lower specific leaf area compared with the other seven species. Finally, one species, Hakea macrocarpa A.Cunn ex.R.Br., had traits that may allow it to tolerate seasonal dryness (through possession of small specific leaf area and cavitation resistant xylem) despite exhibiting cellular water relations that were similar to groundwater-dependent species. We conclude that traits related to water transport and leaf water status differ across species that experience differences in soil water availability and that this enables a diversity of species to exist in this low rainfall environment.

Drought rapidly diminishes the large net CO2 uptake in 2011 over semi-arid Australia.

Each year, terrestrial ecosystems absorb more than a quarter of the anthropogenic carbon emissions, termed as land carbon sink. An exceptionally large land carbon sink anomaly was recorded in 2011, of which more than half was attributed to Australia. However, the persistence and spatially attribution of this carbon sink remain largely unknown. Here we conducted an observation-based study to characterize the Australian land carbon sink through the novel coupling of satellite retrievals of atmospheric CO2 and photosynthesis and in-situ flux tower measures. We show the 2010-11 carbon sink was primarily ascribed to savannas and grasslands. When all biomes were normalized by rainfall, shrublands however, were most efficient in absorbing carbon. We found the 2010-11 net CO2 uptake was highly transient with rapid dissipation through drought. The size of the 2010-11 carbon sink over Australia (0.97 Pg) was reduced to 0.48 Pg in 2011-12, and was nearly eliminated in 2012-13 (0.08 Pg). We further report evidence of an earlier 2000-01 large net CO2 uptake, demonstrating a repetitive nature of this land carbon sink. Given a significant increasing trend in extreme wet year precipitation over Australia, we suggest that carbon sink episodes will exert greater future impacts on global carbon cycle.

The Australian SuperSite Network: A continental, long-term terrestrial ecosystem observatory.

Ecosystem monitoring networks aim to collect data on physical, chemical and biological systems and their interactions that shape the biosphere. Here we introduce the Australian SuperSite Network that, along with complementary facilities of Australia's Terrestrial Ecosystem Research Network (TERN), delivers field infrastructure and diverse, ecosystem-related datasets for use by researchers, educators and policy makers. The SuperSite Network uses infrastructure replicated across research sites in different biomes, to allow comparisons across ecosystems and improve scalability of findings to regional, continental and global scales. This conforms with the approaches of other ecosystem monitoring networks such as Critical Zone Observatories, the U.S. National Ecological Observatory Network; Analysis and Experimentation on Ecosystems, Europe; Chinese Ecosystem Research Network; International Long Term Ecological Research network and the United States Long Term Ecological Research Network. The Australian SuperSite Network currently involves 10 SuperSites across a diverse range of biomes, including tropical rainforest, grassland and savanna; wet and dry sclerophyll forest and woodland; and semi-arid grassland, woodland and savanna. The focus of the SuperSite Network is on using vegetation, faunal and biophysical monitoring to develop a process-based understanding of ecosystem function and change in Australian biomes; and to link this with data streams provided by the series of flux towers across the network. The Australian SuperSite Network is also intended to support a range of auxiliary researchers who contribute to the growing body of knowledge within and across the SuperSite Network, public outreach and education to promote environmental awareness and the role of ecosystem monitoring in the management of Australian environments.

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.

The importance of interacting climate modes on Australia's contribution to global carbon cycle extremes.

The global carbon cycle is highly sensitive to climate-driven fluctuations of precipitation, especially in the Southern Hemisphere. This was clearly manifested by a 20% increase of the global terrestrial C sink in 2011 during the strongest sustained La Niña since 1917. However, inconsistencies exist between El Niño/La Niña (ENSO) cycles and precipitation in the historical record; for example, significant ENSO-precipitation correlations were present in only 31% of the last 100 years, and often absent in wet years. To resolve these inconsistencies, we used an advanced temporal scaling method for identifying interactions amongst three key climate modes (El Niño, the Indian Ocean dipole, and the southern annular mode). When these climate modes synchronised (1999-2012), drought and extreme precipitation were observed across Australia. The interaction amongst these climate modes, more than the effect of any single mode, was associated with large fluctuations in precipitation and productivity. The long-term exposure of vegetation to this arid environment has favoured a resilient flora capable of large fluctuations in photosynthetic productivity and explains why Australia was a major contributor not only to the 2011 global C sink anomaly but also to global reductions in photosynthetic C uptake during the previous decade of drought.

Co-ordination among leaf water relations and xylem vulnerability to embolism of Eucalyptus trees growing along a depth-to-groundwater gradient.

The importance of groundwater resources in arid and semi-arid areas for plant survival is well documented. However, there have been few studies examining the importance and impacts of groundwater availability in mesic environments. The aim of this study was to determine how depth-to-groundwater (DGW) impacts on leaf water relations, leaf structure and branch xylem vulnerability to embolism in a mesic environment. We hypothesize that increasing DGW results in increased resistance to drought stress and that this will be manifested across leaf and branch attributes pertaining to water relations. We further investigate whether there is co-ordination across leaf and branch-scale level responses to increased DGW. Four species were used in this study: Eucalyptus globoidea Blakely, E. piperita Sm., E. sclerophylla (Blakely) L.A.S.Johnson & Blaxell and E. sieberi L.A.S.Johnson. Six sites were chosen along an 11 km transect to span a range of average DGW: 2.4, 4.3, 9.8, 13, 16.3 and 37.5 m. Leaf water relations of trees showed less sensitivity to drought stress as DGW increased. This was reflected in significantly lower leaf turgor loss point and maximum osmotic potential, increased maximum turgor and a reduced leaf relative water content as DGW increased. At shallow DGW sites, minimum diurnal leaf water potentials were generally more negative than leaf water potential at zero turgor, but the reverse was observed at deep sites, indicating a larger growth potential safety margin at deep sites compared with shallow sites. Leaf cell wall elasticity varied independently of DGW. Xylem vulnerability to embolism was quantified as the water potential associated with 50% loss of conductance (P 50). In both summer and winter P 50 was significantly and negatively correlated with DGW. Co-ordination between leaf- and branch-level responses to increase in DGW was apparent, which strongly supports the conclusion that groundwater supply influenced woodland structure and functional behaviour.

Global change-type drought-induced tree mortality: vapor pressure deficit is more important than temperature per se in causing decline in tree health.

Drought-induced tree mortality is occurring across all forested continents and is expected to increase worldwide during the coming century. Regional-scale forest die-off influences terrestrial albedo, carbon and water budgets, and land-surface energy partitioning. Although increased temperatures during drought are widely identified as a critical contributor to exacerbated tree mortality associated with "global-change-type drought", corresponding changes in vapor pressure deficit (D) have rarely been considered explicitly and have not been disaggregated from that of temperature per se. Here, we apply a detailed mechanistic soil-plant-atmosphere model to examine the impacts of drought, increased air temperature (+2°C or +5°C), and increased vapor pressure deficit (D; +1 kPa or +2.5 kPa), singly and in combination, on net primary productivity (NPP) and transpiration and forest responses, especially soil moisture content, leaf water potential, and stomatal conductance. We show that increased D exerts a larger detrimental effect on transpiration and NPP, than increased temperature alone, with or without the imposition of a 3-month drought. Combined with drought, the effect of increased D on NPP was substantially larger than that of drought plus increased temperature. Thus, the number of days when NPP was zero across the 2-year simulation was 13 or 14 days in the control and increased temperature scenarios, but increased to approximately 200 days when D was increased. Drought alone increased the number of days of zero NPP to 88, but drought plus increased temperature did not increase the number of days. In contrast, drought and increased D resulted in the number of days when NPP = 0 increasing to 235 (+1 kPa) or 304 days (+2.5 kPa). We conclude that correct identification of the causes of global change-type mortality events requires explicit consideration of the influence of D as well as its interaction with drought and temperature. This study disaggregates the influence of temperature and vapour pressure deficit on net primary productivity of an Australian woodland and their interactions with drought as potential causal agents in recent widespread forest mortality.

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.

Control of Tamarix in the Western United States: implications for water salvage, wildlife use, and riparian restoration.

Non-native shrub species in the genus Tamarix (saltcedar, tamarisk) have colonized hundreds of thousands of hectares of floodplains, reservoir margins, and other wetlands in western North America. Many resource managers seek to reduce saltcedar abundance and control its spread to increase the flow of water in streams that might otherwise be lost to evapotranspiration, to restore native riparian (streamside) vegetation, and to improve wildlife habitat. However, increased water yield might not always occur and has been substantially lower than expected in water salvage experiments, the potential for successful revegetation is variable, and not all wildlife taxa clearly prefer native plant habitats over saltcedar. As a result, there is considerable debate surrounding saltcedar control efforts. We review the literature on saltcedar control, water use, wildlife use, and riparian restoration to provide resource managers, researchers, and policy-makers with a balanced summary of the state of the science. To best ensure that the desired outcomes of removal programs are met, scientists and resource managers should use existing information and methodologies to carefully select and prioritize sites for removal, apply the most appropriate and cost-effective control methods, and then rigorously monitor control efficacy, revegetation success, water yield changes, and wildlife use.

Invasive capacity of Tamarix ramosissima in a Mojave Desert floodplain: the role of drought.

Tamarix ramosissima (Tamaricaceae) is a woody phreatophyte that has invaded thousands of hectares of floodplain habitat in the southwestern U.S. In this study, we examined the response of gas exchange and stem sap flow of Tamarix and three co-occurring native phreatophytes (Pluchea sericea (Asteraceae), Prosopis pubescens (Fabaceae) and Salix exigua (Salicaceae)) to drought conditions in an early successional floodplain community in the Mojave Desert of southern Nevada. In an analysis of a size/age series of each species across the whole floodplain (both mature and successional stands), stem growth rate was lowest for Tamarix. However, along the same successional chronosequence, Tamarix came to dominate the 50+ year old stands with dense thickets of high stem density. Xylem sap flow, when expressed on a sapwood area basis, was highest in Tamarix under early drought conditions, but comparable between the four species toward the end of the summer dry season. Multivariate analysis of the gas exchange data indicated that the four species differentiated based on water use under early drought conditions and separated based on plant water potential and leaf temperature (indices of drought effects) at the end of the summer dry season. This analysis suggests that the invasive Tamarix is the most drought tolerant of the four species, whereas Salix transpires the most water per unit leaf surface area and is the least tolerant of seasonal water stress. Therefore, Salix appears to be well adapted to early successional communities. However, as floodplains in this arid region become more desiccated with age, Tamarix assumes greater dominance due to its superior drought tolerance relative to native phreatophytes and its ability to produce high density stands and high leaf area.