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Aim: Increased tree mortality linked to droughts and fires is occurring across temperate regions globally. Vegetation recovery has been widely reported; however, less is known about how disturbance may alter forests structurally and functionally across environmental gradients. We examined whether dry forests growing on low‐fertility soils were more resilient to coupled extreme drought and severe fire owing to lower tree mortality rates, higher resprouting success and persistence of juveniles relative to wetter forests on more fertile soils. Location: Fire‐tolerant eucalypt forests of temperate southeastern Australia. Time period: 2020–2023. Major taxa studied: Eucalyptus, Corymbia, Angophora. Methods: Demographic surveys of tree mortality and regeneration in all combinations of dry/wet forest, fertile/less fertile substrates exposed to extreme drought and fire were conducted. We used Bayesian regression modelling to compare tree mortality, diameter, response traits, population structure and occurrence of fire scars between substrates/forest types. Results: Overall mortality (20%–33%) and topkill (34%–41%) were within historically reported ranges for various forests and soil types. However, we observed an atypical trend of increased mortality and topkill in the largest trees, particularly when they had structural damage from past fires. Trees in wet forests on more fertile soils had the highest levels of mortality. Numbers of persistent resprouting juveniles were highest in dry forests on low‐fertility soils. Dry forests growing on low‐fertility soils appear more resilient to compound disturbances due to lower rates of mortality and higher rates of juvenile persistence. Wet forests on more fertile soils may experience greater demographic change due to higher mortality of small and large trees. Main conclusions: Mesic forests on relatively fertile soils were found to be at relatively high risk of demographic change from compound disturbances. Combined, fire and drought are likely to reduce the number of large trees in affected areas, with consequences for forest carbon cycling and storage. [ABSTRACT FROM AUTHOR]

High air temperatures increase atmospheric vapor pressure deficit (VPD) and the severity of drought, threatening forests worldwide. Plants regulate stomata to maximize carbon gain and minimize water loss, resulting in a close coupling between net photosynthesis (Anet) and stomatal conductance (gs). However, evidence for decoupling of gs from Anet under extreme heat has been found. Such a response both enhances survival of leaves during heat events but also quickly depletes available water. To understand the prevalence and significance of this decoupling, we measured leaf gas exchange in 26 tree and shrub species growing in the glasshouse or at an urban site in Sydney, Australia on hot days (maximum Tair > 40°C). We hypothesized that on hot days plants with ample water access would exhibit reduced Anet and use transpirational cooling leading to stomatal decoupling, whereas plants with limited water access would rely on other mechanisms to avoid lethal temperatures. Instead, evidence for stomatal decoupling was found regardless of plant water access. Transpiration of well‐watered plants was 23% higher than model predictions during heatwaves, which effectively cooled leaves below air temperature. For hotter, droughted plants, the increase in transpiration during heatwaves was even more pronounced—gs was 77% higher than model predictions. Stomatal decoupling was found for most broadleaf evergreen and broadleaf deciduous species at the urban site, including some wilted trees with limited water access. Decoupling may simply be a passive consequence of the physical effects of high temperature on plant leaves through increased cuticular conductance of water vapor, or stomatal decoupling may be an adaptive response that is actively regulated by stomatal opening under high temperatures. This temperature response is not yet included in any land surface model, suggesting that model predictions of evapotranspiration may be underpredicted at high temperature and high VPD. [ABSTRACT FROM AUTHOR]

Much attention has been given to the disproportionate streamflow deficits (relative to rainfall deficits) experienced by many catchments during the Millennium Drought (1998–2009) in southeastern Australia, along with lack of post‐drought streamflow recovery in some cases. However, mechanisms behind the coupled hydrologic and ecosystem dynamics are poorly understood. We applied a process‐based ecohydrologic model (RHESSys) in a Melbourne water supply catchment to examine changes in ecohydrologic behavior during and after the drought. Our simulations suggested that average transpiration (green water) was maintained under drought despite a substantial (12%) decrease in average rainfall, meaning that the entire rainfall deficit translated to reduced streamflow (blue water). Altered spatial patterns of vegetation behavior across the terrain helped the ecosystem maintain this unexpectedly high green water use. Decreased transpiration upland was compensated by increases in the riparian zone, which was less water limited and therefore able to meet higher water demand during drought. In the post‐drought period, we found greater transpiration and reduced subsurface water storage relative to pre‐drought, suggesting a longer‐term persistence in altered water partitioning. The post‐drought outcome was attributed to a combination of warmer climate and the persisting effects of the drought on nutrient availability. Given the importance of shifting ecohydrologic patterns across space, our results raise concerns for applying lumped conceptual hydrologic models under nonstationary or extreme conditions. Additionally, the processes we identified have important implications for water supply in Australia's second largest city under projected drying. Plain Language Summary: The Millennium Drought led to unexpectedly large streamflow decreases in catchments across southeastern Australia. This is concerning for water security because climate models suggest the region could experience long‐term rainfall decreases. Using a model that simulates water flow through the landscape, vegetation growth, and nutrient cycling, we investigated drought impacts in a Melbourne water supply catchment. We found that the decrease in rainfall had little impact on catchment‐averaged vegetation water use, meaning that the entire reduction in rainfall was expressed through reduced streamflow (leading to a large proportional decrease). Vegetation water use decreased in most parts of the catchment when rainfall decreased, as would be expected, but this was compensated by increases in the lowland areas where water was less limiting. Water subsidies from upslope allowed these lowland areas to respond to increased water demand during drought. After the drought, the climate was warmer than before the drought and nutrient availability was higher due to a combination of faster decomposition and reduced uptake by vegetation during the drought. Therefore, plant productivity and water use were higher post‐drought than pre‐drought. Our results suggest mechanisms via which water outflow from catchments could be severely reduced under future drying scenarios. Key Points: We used an ecohydrologic model to simulate a Victorian Mountain Ash catchment before, during and after the Millennium DroughtCatchment‐wide transpiration during the drought was maintained at pre‐drought levels via altered spatial patterns of vegetation water usePost‐drought, a combination of warmer temperatures and persisting effects of the drought on nutrients led to even higher transpiration [ABSTRACT FROM AUTHOR]

Drought‐induced vegetation declines have been reported across the globe and may have widespread implications for ecosystem composition, structure, and functions. Thus, it is critical to maximizing our understanding of how vegetation has responded to recent drought extremes. To date, most drought assessments emphasized the importance of drought intensity for vegetation responses. However, drought timing, duration, and repeat exposure all may be important aspects of ecosystem response with the potential for non‐linear effects. Cumulative effects are one such phenomenon, representing the additional decline due to repeated exposure to drought, and indicating gradual loss of ecosystem resistance. This study quantifies the frequency and magnitude of cumulative effects among Australian ecosystems as they responded to the Millennium Drought. Three distinct biophysical variables derived from satellite remote sensing were analyzed, including fraction of photosynthetically absorbed radiation, photosynthetic vegetation cover, and canopy density derived from passive microwave data. Cumulative effects were detected in only 8%–20% of the fire‐free landscape exposed to repeat or long‐duration drought, and could be a statistical artifact. In those limited cases, they approximately doubled drought impacts on leaf abundance, canopy cover, and vegetation density. Cultivated lands and grasslands were the most susceptible to cumulative effects, losing resistance to recurrent droughts, but could be false discovery. Despite being relatively infrequent in forests and savannas, cumulative effects caused larger additional declines in these ecosystems. Overall, our study demonstrates that repeated exposure appears to have limited influence on the magnitude of drought impacts on canopy structure affecting only a few areas. Key Points: Cumulative effects affected 8%∼20% of fire‐free landscape exposed to repeat or long‐duration droughts during the Millennium Drought periodDespite being relatively infrequent, cumulative effects doubled drought impacts in areas where they occurredLarge effects in forests and savannas implies a greater risk of carbon release given recurrent drought events [ABSTRACT FROM AUTHOR]

The Millennium Drought lasted more than a decade and is notable for causing persistent shifts in the relationship between rainfall and runoff in many southeastern Australian catchments. Research to date has successfully characterised where and when shifts occurred and explored relationships with potential drivers, but a convincing physical explanation for observed changes in catchment behaviour is still lacking. Originating from a large multi-disciplinary workshop, this paper presents and evaluates a range of hypothesised process explanations of flow response to the Millennium Drought. The hypotheses consider climatic forcing, vegetation, soil moisture dynamics, groundwater, and anthropogenic influence. The hypotheses are assessed against evidence both temporally (e.g. why was the Millennium Drought different to previous droughts?) and spatially (e.g. why did rainfall–runoff relationships shift in some catchments but not in others?). Thus, the strength of this work is a large-scale assessment of hydrologic changes and potential drivers. Of 24 hypotheses, 3 are considered plausible, 10 are considered inconsistent with evidence, and 11 are in a category in between, whereby they are plausible yet with reservations (e.g. applicable in some catchments but not others). The results point to the unprecedented length of the drought as the primary climatic driver, paired with interrelated groundwater processes, including declines in groundwater storage, altered recharge associated with vadose zone expansion, and reduced connection between subsurface and surface water processes. Other causes include increased evaporative demand and harvesting of runoff by small private dams. Finally, we discuss the need for long-term field monitoring, particularly targeting internal catchment processes and subsurface dynamics. We recommend continued investment in the understanding of hydrological shifts, particularly given their relevance to water planning under climate variability and change. [ABSTRACT FROM AUTHOR]

In 2019, south-eastern Australia experienced its driest and hottest year on record, resulting in massive canopy dieback events in eucalypt dominated forests. A subsequent period of high precipitation in 2020 provided a rare opportunity to quantify the impacts of extreme drought and consequent recovery. We quantified canopy health and hydraulic impairment (native percent loss of hydraulic conductivity, PLC) of 18 native tree species growing at 15 sites that were heavily impacted by the drought both during and 8–10 months after the drought. Most species exhibited high PLC during drought (PLC:65.1 ± 3.3%), with no clear patterns across sites or species. Heavily impaired trees (PLC > 70%) showed extensive canopy browning. In the post-drought period, most surviving trees exhibited hydraulic recovery (PLC:26.1 ± 5.1%), although PLC remained high in some trees (50–70%). Regained hydraulic function (PLC < 50%) corresponded to decreased canopy browning indicating improved tree health. Similar drought (37.1 ± 4.2%) and post-drought (35.1 ± 4.4%) percentages of basal area with dead canopy suggested that trees with severely compromised canopies immediately after drought were not able to recover. This dataset provides insights into the impacts of severe natural drought on the health of mature trees, where hydraulic failure is a major contributor in canopy dieback and tree mortality during extreme drought events. [ABSTRACT FROM AUTHOR]

Aim: To estimate loss of above‐ground carbon (AGC) and conversion of live carbon to dead carbon following understorey and canopy fire. Location: South‐eastern Australia. Time period: 2019–2020. Major taxa studied: Four widespread resprouting eucalypt forests. Methods: Above‐ground carbon was measured in 15 plots in each of four forest types one‐year post‐fire. We also assessed topkill, that is, trees subject to canopy loss that failed to resprout epicormically. Results: While canopy fire was associated with greater declines in AGC than understorey fire, this was only statistically significant for only one forest type, where AGC declined from 154 to 85 Mg C ha−1 following canopy fire. Significant post‐fire increases in dead AGC were observed in one forest type, where dead carbon increased from 22 to 60% after canopy fire. Topkill of trees following canopy fire (48–78% of stems) was higher than topkill after understorey fire (36–53% of stems) and in unburnt forest (12–55%). Topkill occurred primarily in small‐diameter stems. Consequently, there was no effect of fire on the proportion of dead AGC in trees, with the exception of the forest with lowest productivity (i.e., lowest biomass) and lowest annual rainfall, where dead tree carbon increased from 8% in unburnt forest to 13 and 53% after understorey and canopy fire, respectively. AGC in understorey vegetation and coarse woody debris was similar or lower in burnt compared with unburnt forest. Litter carbon was significantly lower and pyrogenic carbon significantly higher in burnt forest, with no difference between understorey and canopy fire. Main conclusions While increased fire severity was associated with increased changes to carbon stocks, there were differences among forest types. Specifically, the driest forest type had the highest rates of topkill following canopy fire. These results highlight the importance of spatial variability in fire severity and forest type in determining the effects of fire on carbon stocks. [ABSTRACT FROM AUTHOR]

Summary: Predicting species‐level responses to drought at the landscape scale is critical to reducing uncertainty in future terrestrial carbon and water cycle projections.We embedded a stomatal optimisation model in the Community Atmosphere Biosphere Land Exchange (CABLE) land surface model and parameterised the model for 15 canopy dominant eucalypt tree species across South‐Eastern Australia (mean annual precipitation range: 344–1424 mm yr−1). We conducted three experiments: applying CABLE to the 2017–2019 drought; a 20% drier drought; and a 20% drier drought with a doubling of atmospheric carbon dioxide (CO2).The severity of the drought was highlighted as for at least 25% of their distribution ranges, 60% of species experienced leaf water potentials beyond the water potential at which 50% of hydraulic conductivity is lost due to embolism. We identified areas of severe hydraulic stress within‐species' ranges, but we also pinpointed resilience in species found in predominantly semiarid areas. The importance of the role of CO2 in ameliorating drought stress was consistent across species.Our results represent an important advance in our capacity to forecast the resilience of individual tree species, providing an evidence base for decision‐making around the resilience of restoration plantings or net‐zero emission strategies. [ABSTRACT FROM AUTHOR]

Between late 2015 and early 2016, more than 7,000 ha of mangrove forest died along the coastline of the Gulf of Carpentaria, in northern Australia. This massive die-off was preceded by a strong 2015/2016 El Niño event, resulting in lower precipitation, a drop in sea level and higher than average temperatures in northern Australia. In this study, we investigated the role of hydraulic failure in the mortality and recovery of the dominant species, Avicennia marina , 2 years after the mortality event. We measured predawn water potential (Ψpd) and percent loss of stem hydraulic conductivity (PLC) in surviving individuals across a gradient of impact. We also assessed the vulnerability to drought-induced embolism (Ψ50) for the species. Areas with severe canopy dieback had higher native PLC (39%) than minimally impacted areas (6%), suggesting that hydraulic recovery was ongoing. The high resistance of A. marina to water-stress-induced embolism (Ψ50 = −9.6 MPa), indicates that severe water stress (Ψpd < −10 MPa) would have been required to cause mortality in this species. Our data indicate that the natural gradient of water-stress enhanced the impact of El Niño, leading to hydraulic failure and mortality in A. marina growing on severely impacted (SI) zones. It is likely that lowered sea levels and less frequent inundation by seawater, combined with lower inputs of fresh water, high evaporative demand and high temperatures, led to the development of hyper-salinity and extreme water stress during the 2015/16 summer. [ABSTRACT FROM AUTHOR]

Climate change is projected to increase the imbalance between the supply (precipitation) and atmospheric demand for water (i.e., increased potential evapotranspiration), stressing plants in water-limited environments. Plants may be able to offset increasing aridity because rising CO2 increases water use efficiency. CO2 fertilization has also been cited as one of the drivers of the widespread "greening" phenomenon. However, attributing the size of this CO2 fertilization effect is complicated, due in part to a lack of long-term vegetation monitoring and interannual- to decadal-scale climate variability. In this study we asked the question of how much CO2 has contributed towards greening. We focused our analysis on a broad aridity gradient spanning eastern Australia's woody ecosystems. Next we analyzed 38 years of satellite remote sensing estimates of vegetation greenness (normalized difference vegetation index, NDVI) to examine the role of CO2 in ameliorating climate change impacts. Multiple statistical techniques were applied to separate the CO2 -attributable effects on greening from the changes in water supply and atmospheric aridity. Widespread vegetation greening occurred despite a warming climate, increases in vapor pressure deficit, and repeated record-breaking droughts and heat waves. Between 1982–2019 we found that NDVI increased (median 11.3 %) across 90.5 % of the woody regions. After masking disturbance effects (e.g., fire), we statistically estimated an 11.7 % increase in NDVI attributable to CO2 , broadly consistent with a hypothesized theoretical expectation of an 8.6 % increase in water use efficiency due to rising CO2. In contrast to reports of a weakening CO2 fertilization effect, we found no consistent temporal change in the CO2 effect. We conclude rising CO2 has mitigated the effects of increasing aridity, repeated record-breaking droughts, and record-breaking heat waves in eastern Australia. However, we were unable to determine whether trees or grasses were the primary beneficiary of the CO2 -induced change in water use efficiency, which has implications for projecting future ecosystem resilience. A more complete understanding of how CO2 -induced changes in water use efficiency affect trees and non-tree vegetation is needed. [ABSTRACT FROM AUTHOR]

Understanding vegetation recovery after drought is critical for projecting vegetation dynamics in future climates. From 1997 to 2009, Australia experienced a long‐lasting drought known as the Millennium Drought (MD), which led to widespread reductions in vegetation productivity. However, vegetation recovery post‐drought and its determinants remain unclear. This study leverages remote sensing products from different sources—fraction of absorbed photosynthetically active radiation (FPAR), based on optical data, and canopy density, derived from microwave data—and random forest algorithms to assess drought recovery over Australian natural vegetation during a 20‐year period centered on the MD. Post‐drought recovery was prevalent across the continent, with 6 out of 10 drought events seeing full recovery within about 6 months. Canopy density was slower to recover than leaf area seen in FPAR. The probability of full recovery was most strongly controlled by drought return interval, post‐drought hydrological condition, and drought length. Full recovery was seldom observed when drought events occurred at intervals of 3 months or less, and moderately dry (standardized water balance anomaly [SWBA] within [−1, −0.76]) post‐drought conditions resulted in less complete recovery than wet (SWBA > 0.3) post‐drought conditions. Press droughts, which are long term but not extreme, delayed recovery more than pulse droughts (short term but extreme) and led to a higher frequency of persistent decline. Following press droughts, the frequency of persistent decline differed little among biome types but peaked in semi‐arid regions across aridity levels. Forests and savanna required the longest recovery times for press drought, while grasslands were the slowest to recover for pulse drought. This study provides quantitative thresholds that could be used to improve the modeling of ecosystem dynamics post‐drought. [ABSTRACT FROM AUTHOR]

Summary: Eastern Australia was subject to its hottest and driest year on record in 2019. This extreme drought resulted in massive canopy die‐back in eucalypt forests. The role of hydraulic failure and tree size on canopy die‐back in three eucalypt tree species during this drought was examined.We measured pre‐dawn and midday leaf water potential (Ψleaf), per cent loss of stem hydraulic conductivity and quantified hydraulic vulnerability to drought‐induced xylem embolism. Tree size and tree health was also surveyed.Trees with most, or all, of their foliage dead exhibited high rates of native embolism (78–100%). This is in contrast to trees with partial canopy die‐back (30–70% canopy die‐back: 72–78% native embolism), or relatively healthy trees (little evidence of canopy die‐back: 25–31% native embolism). Midday Ψleaf was significantly more negative in trees exhibiting partial canopy die‐back (−2.7 to −6.3 MPa), compared with relatively healthy trees (−2.1 to −4.5 MPa). In two of the species the majority of individuals showing complete canopy die‐back were in the small size classes.Our results indicate that hydraulic failure is strongly associated with canopy die‐back during drought in eucalypt forests. Our study provides valuable field data to help constrain models predicting mortality risk. [ABSTRACT FROM AUTHOR]

Land surface models underpin coupled climate model projections of droughts and heatwaves. However, the lack of simultaneous observations of individual components of evapotranspiration, concurrent with root-zone soil moisture, has limited previous model evaluations. Here, we use a comprehensive set of observations from a water-limited site in southeastern Australia including both evapotranspiration and soil moisture to a depth of 4.5 m to evaluate the Community Atmosphere-Biosphere Land Exchange (CABLE) land surface model. We demonstrate that alternative process representations within CABLE had the capacity to improve simulated evapotranspiration, but not necessarily soil moisture dynamics–highlighting problems of model evaluations against water fluxes alone. Our best simulation was achieved by resolving a soil evaporation bias, using a more realistic initialisation of the groundwater aquifer state and higher vertical soil resolution informed by observed soil properties, and further calibrating soil hydraulic conductivity. Despite these improvements, the role of the empirical soil moisture stress function in influencing the simulated water fluxes remained important: using a site-calibrated function reduced the soil water stress on plants by 36 % during drought and 23 % at other times. These changes in CABLE not only improve the seasonal cycle of evapotranspiration but also affect the latent and sensible heat fluxes during droughts and heatwaves. The range of parameterisations tested led to differences of ∼150 Wm-2 in the simulated latent heat flux during a heatwave, implying a strong impact of parameterisations on the capacity for evaporative cooling and feedbacks to the boundary layer (when coupled). Overall, our results highlight the opportunity to advance the capability of land surface models to capture water cycle processes, particularly during meteorological extremes, when sufficient observations of both evapotranspiration fluxes and soil moisture profiles are available. [ABSTRACT FROM AUTHOR]

South‐East Australia has recently been subjected to two of the worst droughts in the historical record (Millennium Drought, 2000–2009 and Big Dry, 2017–2019). Unfortunately, a lack of forest monitoring has made it difficult to determine whether widespread tree mortality has resulted from these droughts. Anecdotal observations suggest the Big Dry may have led to more significant tree mortality than the Millennium drought. Critically, to be able to robustly project future expected climate change effects on Australian vegetation, we need to assess the vulnerability of Australian trees to drought. Here we implemented a model of plant hydraulics into the Community Atmosphere Biosphere Land Exchange (CABLE) land surface model. We parameterized the drought response behaviour of five broad vegetation types, based on a common garden dry‐down experiment with species originating across a rainfall gradient (188–1,125 mm/year) across South‐East Australia. The new hydraulics model significantly improved (~35%–45% reduction in root mean square error) CABLE's previous predictions of latent heat fluxes during periods of water stress at two eddy covariance sites in Australia. Landscape‐scale predictions of the greatest percentage loss of hydraulic conductivity (PLC) of about 40%–60%, were broadly consistent with satellite estimates of regions of the greatest change in both droughts. In neither drought did CABLE predict that trees would have reached critical PLC in widespread areas (i.e. it projected a low mortality risk), although the model highlighted critical levels near the desert regions of South‐East Australia where few trees live. Overall, our experimentally constrained model results imply significant resilience to drought conferred by hydraulic function, but also highlight critical data and scientific gaps. Our approach presents a promising avenue to integrate experimental data and make regional‐scale predictions of potential drought‐induced hydraulic failure. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The iso/anisohydric continuum describes how plants regulate leaf water potential and is commonly used to classify species drought response strategies. However, drought response strategies comprise more than just this continuum, incorporating a suite of stomatal and hydraulic traits.Using a common garden experiment, we compared and contrasted four metrics commonly used to describe water use strategy during drought in 10 eucalyptus species comprising four major ecosystems in eastern Australia. We examined the degree to which these metrics were aligned with key stomatal and hydraulic traits related to plant water use and drought tolerance.Species rankings of water use strategy were inconsistent across four metrics. A newer metric (Hydroscape) was strongly linked to various plant traits, including the leaf turgor loss (TLP), water potential at stomatal closure (Pgs90), leaf and stem hydraulic vulnerability to embolism (PL50 and Px50), safety margin of hydraulic segmentation (HSMHS), maximum stomatal conductance (gsmax) and Huber value (HV). In addition, Hydroscape was correlated with climatic variables representing the water availability at the seed source site.Along the continuum of water regulation strategy, species with narrow Hydroscapes tended to occupy mesic regions and exhibit high TLP, PL50 and Px50 values and narrow HSMHS. High gsmax recorded in species with broad hydroscapes was also associated with high HV.Despite a fourfold difference in Hydroscape area, all species closed their stomata prior to the onset of hydraulic dysfunction, suggesting a common stomatal response across species that minimizes embolism risk during drought. Hydroscape area is useful in bridging stomatal regulation, hydraulic architecture and species drought tolerance, thus providing insight into species water use strategies. A plain language summary is available for this article. [ABSTRACT FROM AUTHOR]

Recent experimental evidence suggests that during heat extremes, wooded ecosystems may decouple photosynthesis and transpiration, reducing photosynthesis to near zero but increasing transpiration into the boundary layer. This feedback may act to dampen, rather than amplify, heat extremes in wooded ecosystems. We examined eddy covariance databases (OzFlux and FLUXNET2015) to identify whether there was field-based evidence to support these experimental findings. We focused on two types of heat extremes: (i) the 3 days leading up to a temperature extreme, defined as including a daily maximum temperature >37 ∘ C (similar to the widely used TXx metric), and (ii) heatwaves, defined as 3 or more consecutive days above 35 ∘ C. When focusing on (i), we found some evidence of reduced photosynthesis and sustained or increased latent heat fluxes at seven Australian evergreen wooded flux sites. However, when considering the role of vapour pressure deficit and focusing on (ii), we were unable to conclusively disentangle the decoupling between photosynthesis and latent heat flux from the effect of increasing the vapour pressure deficit. Outside of Australia, the Tier-1 FLUXNET2015 database provided limited scope to tackle this issue as it does not sample sufficient high temperature events with which to probe the physiological response of trees to extreme heat. Thus, further work is required to determine whether this photosynthetic decoupling occurs widely, ideally by matching experimental species with those found at eddy covariance tower sites. Such measurements would allow this decoupling mechanism to be probed experimentally and at the ecosystem scale. Transpiration during heatwaves remains a key issue to resolve, as no land surface model includes a decoupling mechanism, and any potential dampening of the land–atmosphere amplification is thus not included in climate model projections. [ABSTRACT FROM AUTHOR]

We tested the ability of a model to scale gas exchange from leaf level to whole-tree level by: (1) measuring leaf gas exchange in the canopy of 10 trees in a tall Eucalyptus delegatensis RT Baker forest in NSW, Australia; (2) monitoring sap flow of the same 10 trees during the measurement week; and (3) using an individual-tree-based model (MAESTRA) to link the two sets of measurements. Photosynthesis and stomatal conductance components of the model were parameterized with the leaf gas exchange data, and canopy structure was parameterized with crown heights, dimensions and leaf areas of each of the measurement trees and up to 45 neighboring trees. Transpiration of the measurement trees was predicted by the model and compared with sap flow data. Leaf gas exchange parameters were similar for all 10 trees, with the exception of two smaller trees that had relatively low stomatal conductances. We hypothesize that these trees may have experienced water stress as a result of competition from large neighboring trees. The model performed well, and in most cases, was able to replicate the time course of tree transpiration. Maximum rates of transpiration were higher than measured rates for some trees and lower than measured rates for others, which may have been a result of inaccuracy in estimating tree leaf area. There was a small lag (about 15-30 minutes) between sap flow and modeled transpiration for some trees in the morning, likely associated with use of water stored in stems. The model also captured patterns of variation in sap flow among trees. Overall, the study confirms the ability of models to estimate forest canopy transpiration from leaf-level measurements. [ABSTRACT FROM AUTHOR]

Globally, fire regimes are being altered by changing climatic conditions. New fire regimes have the potential to drive species extinctions and cause ecosystem state changes, with a range of consequences for ecosystem services. Despite the co-occurrence of forest fires with drought, current approaches to modelling flammability largely overlook the large body of research into plant vulnerability to drought. Here, we outline the mechanisms through which plant responses to drought may affect forest flammability, specifically fuel moisture and the ratio of dead to live fuels. We present a framework for modelling live fuel moisture content (moisture content of foliage and twigs) from soil water content and plant traits, including rooting patterns and leaf traits such as the turgor loss point, osmotic potential, elasticity and leaf mass per area. We also present evidence that physiological drought stress may contribute to previously observed fuel moisture thresholds in south-eastern Australia. Of particular relevance is leaf cavitation and subsequent shedding, which transforms live fuels into dead fuels, which are drier, and thus easier to ignite. We suggest that capitalising on drought research to inform wildfire research presents a major opportunity to develop new insights into wildfires, and new predictive models of seasonal fuel dynamics. [ABSTRACT FROM AUTHOR]

A commentary to an article about net primary production (NPP) to gross primary production (GPP) is presented. In the article, researchers examined the annual carbon budgets for forest location across the U.S., Australia and New Zealand. The authors offer an explanation on how to calculate GPP. They emphasize that they do not challenge the hypothesis that NPP/GPP is constant for forests.

Abstract: Resolving ecophysiological processes in elevated atmospheric CO2 (C a) at scales larger than single leaves poses significant challenges. Here, we describe a field-based experimental system designed to grow trees up to 9m tall in elevated C a with the capacity to control air temperature and simultaneously measure whole-tree gas exchange. In western Sydney, Australia, we established the Hawkesbury Forest Experiment (HFE) where we built whole-tree chambers (WTC) to measure whole-tree CO2 and water fluxes of an evergreen broadleaf tree, Eucalyptus saligna. A single E. saligna tree was grown from seedling to small tree within each of 12 WTCs; six WTCs were maintained at ambient C a and six WTCs were maintained at elevated C a, targeted at ambient C a +240μmolmol−1. All 12 WTCs were controlled to track ambient outside air temperature (T air) and air water vapour deficit (D air). During the experimental period, T air, D air and C a in the WTCs were within 0.5°C, 0.3kPa, and 15μmolmol−1 of the set-points for 90% of the time, respectively. Diurnal responses of whole-tree CO2 and water vapour fluxes are analysed, demonstrating the ability of the tree chamber system to measure rapid environmental responses of these fluxes of entire trees. The light response of CO2 uptake for entire trees showed a clear diurnal hysteresis, attributed to stomatal closure at high D air. Tree scale CO2 fluxes confirm the hypothesised deleterious effect of chilling night-time temperatures on whole-tree carbon gain in this subtropical Eucalyptus. The whole-tree chamber flux data add an invaluable scale to measurements in both ambient and elevated C a and allow us to elucidate the mechanisms driving tree productivity responses to elevated C a in interaction with water availability and temperature. [Copyright &y& Elsevier]

• We examined the spatiotemporal drivers of live fuel moisture content (LFMC). • Models developed from pressure-volume curve data predicted seasonal LFMC. • Leaf dry mass traits predicted maximum LFMC across a large climatic gradient. • Maximum LFMC was correlated with aridity, with lower values in arid sites. • Including leaf traits in LFMC models is likely to improve their performance. Large forest fires generally occur when the moisture content of fuels is low. For live fuels, our understanding of the physiological basis of variation in moisture content has recently advanced. However, process-based models of live fuel moisture content (LFMC) remain elusive. Here, we aim to further our understanding of the role of physiological mechanisms and plant functional traits in driving spatiotemporal variations in LFMC. We examined whether temporal variation in LFMC could be predicted from pressure-volume curve data, which measures leaf water potential and water content on cut shoots dehydrating on a bench. We also examined whether leaf dry mass traits could predict spatial variation in maximum LFMC. We undertook our study in eucalypt forests and woodlands spanning a large climatic gradient in eastern Australia. We found that LFMC models developed from pressure-volume curves reliably predicted seasonal variation in LFMC across four co-occurring species. A two-phase LFMC model, which fit models above and below the turgor loss point (mean absolute error = 3.7-33.2%), performed similarly well to a simple linear model (mean absolute error = 3.4-35.3%). Across a large climatic gradient, the maximum LFMC of 16 species was correlated with specific leaf area (R 2 = 0.54), with the exception of one species with terete terminal stems. Maximum LFMC was highly correlated with aridity (R 2 = 0.82), with lower LFMC observed in more arid sites. Our study demonstrates that spatiotemporal dynamics of LFMC are governed by both leaf dry mass traits and the relationship between leaf water potential and water content, which in turn is determined by traits such as cell wall elasticity. Thus, incorporating these traits into models of LFMC, whether these models are based on drought indices, soil moisture, or remotely sensed imagery, is likely to improve overall model performance, and subsequently improve forecasts of wildfire danger. [ABSTRACT FROM AUTHOR]

In 2020, the Australian and New Zealand flux research and monitoring network, OzFlux, celebrated its 20 th anniversary by reflecting on the lessons learned through two decades of ecosystem studies on global change biology. OzFlux is a network not only for ecosystem researchers, but also for those 'next users' of the knowledge, information and data that such networks provide. Here, we focus on eight lessons across topics of climate change and variability, disturbance and resilience, drought and heat stress and synergies with remote sensing and modelling. In distilling the key lessons learned, we also identify where further research is needed to fill knowledge gaps and improve the utility and relevance of the outputs from OzFlux. Extreme climate variability across Australia and New Zealand (droughts and flooding rains) provides a natural laboratory for a global understanding of ecosystems in this time of accelerating climate change. As evidence of worsening global fire risk emerges, the natural ability of these ecosystems to recover from disturbances, such as fire and cyclones, provides lessons on adaptation and resilience to disturbance. Drought and heatwaves are common occurrences across large parts of the region and can tip an ecosystem's carbon budget from a net CO 2 sink to a net CO 2 source. Despite such responses to stress, ecosystems at OzFlux sites show their resilience to climate variability by rapidly pivoting back to a strong carbon sink upon the return of favourable conditions. Located in under-represented areas, OzFlux data have the potential for reducing uncertainties in global remote sensing products, and these data provide several opportunities to develop new theories and improve our ecosystem models. The accumulated impacts of these lessons over the last 20 years highlights the value of long-term flux observations for natural and managed systems. A future vision for OzFlux includes ongoing and newly developed synergies with ecophysiologists, ecologists, geologists, remote sensors and modellers., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

While trees can acclimate to warming, there is concern that tropical rainforest species may be less able to acclimate because they have adapted to a relatively stable thermal environment. Here we tested whether the physiological adjustments to warming differed among Australian tropical, subtropical and warm-temperate rainforest trees. Photosynthesis and respiration temperature responses were quantified in six Australian rainforest seedlings of tropical, subtropical and warm-temperate climates grown across four growth temperatures in a glasshouse. Temperature-response models were fitted to identify mechanisms underpinning the response to warming. Tropical and subtropical species had higher temperature optima for photosynthesis (T optA ) than temperate species. There was acclimation of T optA to warmer growth temperatures. The rate of acclimation (0.35-0.78°C °C -1 ) was higher in tropical and subtropical than in warm-temperate trees and attributed to differences in underlying biochemical parameters, particularly increased temperature optima of V cmax25 and J max25 . The temperature sensitivity of respiration (Q 10 ) was 24% lower in tropical and subtropical compared with warm-temperate species. Overall, tropical and subtropical species had a similar capacity to acclimate to changes in growth temperature as warm-temperate species, despite being grown at higher temperatures. Quantifying the physiological acclimation in rainforests can improve accuracy of future climate predictions and assess their potential vulnerability to warming., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

Rising atmospheric [CO 2 ] (C a ) generally enhances tree growth if nutrients are not limiting. However, reduced water availability and elevated evaporative demand may offset such fertilization. Trees with access to deep soil water may be able to mitigate such stresses and respond more positively to C a . Here, we sought to evaluate how increased vapor pressure deficit and reduced precipitation are likely to modify the impact of elevated C a (eC a ) on tree productivity in an Australian Eucalyptus saligna Sm. plantation with access to deep soil water. We parameterized a forest growth simulation model (GOTILWA+) using data from two field experiments on E. saligna: a 2-year whole-tree chamber experiment with factorial C a (ambient =380, elevated =620 μmol mol -1 ) and watering treatments, and a 10-year stand-scale irrigation experiment. Model evaluation showed that GOTILWA+ can capture the responses of canopy C uptake to (1) rising vapor pressure deficit (D) under both C a treatments; (2) alterations in tree water uptake from shallow and deep soil layers during soil dry-down; and (3) the impact of irrigation on tree growth. Simulations suggest that increasing C a up to 700 μmol mol -1 alone would result in a 33% increase in annual gross primary production (GPP) and a 62% increase in biomass over 10 years. However, a combined 48% increase in D and a 20% reduction in precipitation would halve these values. Our simulations identify high D conditions as a key limiting factor for GPP. They also suggest that rising C a will compensate for increasing aridity limitations in E. saligna trees with access to deep soil water under non-nutrient limiting conditions, thereby reducing the negative impacts of global warming upon this eucalypt species. Simulation models not accounting for water sources available to deep-rooting trees are likely to overestimate aridity impacts on forest productivity and C stocks., (© 2021 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Vapour pressure deficit (D) is projected to increase in the future as temperature rises. In response to increased D, stomatal conductance (gs) and photosynthesis (A) are reduced, which may result in significant reductions in terrestrial carbon, water and energy fluxes. It is thus important for gas exchange models to capture the observed responses of gs and A with increasing D. We tested a series of coupled A-gs models against leaf gas exchange measurements from the Cumberland Plain Woodland (Australia), where D regularly exceeds 2 kPa and can reach 8 kPa in summer. Two commonly used A-gs models were not able to capture the observed decrease in A and gs with increasing D at the leaf scale. To explain this decrease in A and gs, two alternative hypotheses were tested: hydraulic limitation (i.e., plants reduce gs and/or A due to insufficient water supply) and non-stomatal limitation (i.e., downregulation of photosynthetic capacity). We found that the model that incorporated a non-stomatal limitation captured the observations with high fidelity and required the fewest number of parameters. Whilst the model incorporating hydraulic limitation captured the observed A and gs, it did so via a physical mechanism that is incorrect. We then incorporated a non-stomatal limitation into the stand model, MAESPA, to examine its impact on canopy transpiration and gross primary production. Accounting for a non-stomatal limitation reduced the predicted transpiration by ~19%, improving the correspondence with sap flow measurements, and gross primary production by ~14%. Given the projected global increases in D associated with future warming, these findings suggest that models may need to incorporate non-stomatal limitation to accurately simulate A and gs in the future with high D. Further data on non-stomatal limitation at high D should be a priority, in order to determine the generality of our results and develop a widely applicable model., (© The Author(s) 2019. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Plants show flexible acclimation of leaf photosynthesis to temperature that depends both on their prevailing growth environment and the climate where they originated. This acclimation has been shown to involve changes in the temperature responses of the apparent maximum rate of Rubisco carboxylation (Vcmax) and apparent maximum rate of electron transport (Jmax), as well as changes in the ratio of these parameters. We asked whether such changes in photosynthetic biochemistry attributable to climate of origin are similar in nature and magnitude to those attributable to growth environment. To address this question, we measured temperature responses of photosynthesis and chlorophyll fluorescence on six Eucalyptus species from diverse geographical and climatic regions growing in a common garden. Measurements were made in three seasons, allowing us to compare interspecific differences with seasonal changes. We found significant interspecific differences in apparent Vcmax and Jmax standardized to 25 °C, but there were no significant differences in the temperature responses of these parameters among species. Comparing data across seasons, we found significant seasonal changes in apparent Vcmax25, but not in Jmax25, causing a change in their ratio (J/V ratio). However, there were no seasonal changes in the temperature response of either parameter. We concluded that the growth environment had a much larger effect on temperature response than climate of origin among this set of species. Mean daytime temperature increased by 15 °C from winter to summer, whereas we estimated that the seasonal change in J/V ratio would cause a change in the optimum temperature (Topt) for gross photosynthesis of 3.6 °C. Use of a general relationship to describe photosynthetic temperature acclimation resulted in a strong underestimation of the Topt for photosynthesis for these species. Our results indicated that variation in photosynthetic temperature responses cannot be captured in one simple relationship with growth temperature. Further comparative research on species groups will be needed to develop a basis for modelling these interspecific differences in plant temperature acclimation.

Models of stomatal conductance (g(s)) are based on coupling between g(s) and CO(2) assimilation (A(net)), and it is often assumed that the slope of this relationship ('g(1) ') is constant across species. However, if different plant species have adapted to different access costs of water, then there will be differences in g(1) among species. We hypothesized that g(1) should vary among species adapted to different climates, and tested the theory and its linkage to plant hydraulics using four Eucalyptus species from different climatic origins in a common garden. Optimal stomatal theory predicts that species from sub-humid zones have a lower marginal water cost of C gain, hence lower g(1) than humid-zone species. In agreement with the theory that g(1) is related to tissue carbon costs for water supply, we found a relationship between wood density and g(1) across Eucalyptus species of contrasting climatic origins. There were significant reductions in the parameter g(1) during drought in humid but not sub-humid species, with the latter group maintaining g(1) in drought. There are strong differences in stomatal behaviour among related tree species in agreement with optimal stomatal theory, and these differences are consistent with the economics involved in water uptake and transport for carbon gain., (© 2012 Blackwell Publishing Ltd.)

The response of photosynthesis to temperature is a central facet of plant response to climate. Such responses have been found to be highly variable among species and among studies. Understanding this variability is key when trying to predict the effects of rising global temperatures on plant productivity. There are three major factors affecting the response of leaf net photosynthesis to temperature (A(n)-T): (i) photosynthetic biochemistry, (ii) respiration and (iii) vapour pressure deficit (D) and stomatal sensitivity to vapour pressure deficit during measurements. The overall goal of our study was to quantify the relative contribution of each of these factors in determining the response of A(n) to temperature. We first conducted a sensitivity analysis with a coupled photosynthesis-stomatal (A(n)-g(s)) model, using ranges for parameters of each factor taken from the literature, and quantified how these parameters affected the A(n)-T response. Second, we applied the A(n)-g(s) model to two example sets of field data, which had different optimum temperatures (T(opt)) of A(n), to analyse which factors were most important in causing the difference. We found that each of the three factors could have an equally large effect on T(opt) of A(n). In our comparison between two field datasets, the major cause for the difference in T(opt) was not the biochemical component, but rather the differences in respiratory components and in D conditions during measurements. We concluded that shifts in A(n)-T responses are not always driven by acclimation of photosynthetic biochemistry, but can result from other factors. The D conditions during measurements and stomatal responses to D also need to be quantified if we are to better understand and predict shifts in A(n)-T with climate.