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Optimal stomatal theory predicts that stomata operate to maximise photosynthesis (Anet) and minimise transpirational water loss to achieve optimal intrinsic water-use efficiency (iWUE). We tested whether this theory can predict stomatal responses to elevated atmospheric CO2 (eCO2), and whether it can capture differences in responsiveness among woody plant functional types (PFTs). We conducted a meta-analysis of tree studies of the effect of eCO2 on iWUE and its components Anet and stomatal conductance (gs). We compared three PFTs, using the unified stomatal optimisation (USO) model to account for confounding effects of leaf–air vapour pressure difference (D). We expected smaller gs, but greater Anet, responses to eCO2 in gymnosperms compared with angiosperm PFTs. We found that iWUE increased in proportion to increasing eCO2 in all PFTs, and that increases in Anet had stronger effects than reductions in gs. The USO model correctly captured stomatal behaviour with eCO2 across most datasets. The chief difference among PFTs was a lower stomatal slope parameter (g1) for the gymnosperm, compared with angiosperm, species. Land surface models can use the USO model to describe stomatal behaviour under changing atmospheric CO2 conditions. AG gratefully acknowledges a studentship provided by the John Horseman Trust and the University of Birmingham. The BIFoR FACE facility is supported by the JABBS Foundation, the University of Birmingham and the John Horseman Trust. ARMK acknowledges support from the UK Natural Environment Research Council through grant NE/S015833/1. MJ and BEM acknowledge funding from the Australian Research Council (DE210101654, FL190100003).

Mistletoes are important co-contributors to tree mortality globally, particularly during droughts. In Australia, mistletoe distributions are expanding in temperate woodlands, while their hosts have experienced unprecedented heat and drought stress in recent years. We investigated whether the excessive water use of mistletoes increased the probability of xylem emboli in a mature woodland during the recent record drought that was compounded by multiple heatwaves. We continuously recorded transpiration ($T_{SLA}$) of infected and uninfected branches from two eucalypt species over two summers, monitored stem and leaf water potentials ($\Psi $) and used hydraulic vulnerability curves to estimate percent loss in conductivity (PLC) for each species. Variations in weather (vapor pressure deficit, photosynthetically active radiation, soil water content), host species and % mistletoe foliage explained 78% of hourly $T_{SLA}$. While mistletoe acted as an uncontrollable sink for water in the host even during typical summer days, daily $T_{SLA}$ increased up to 4-fold in infected branches on hot days, highlighting the previously overlooked importance of temperature stress in amplifying water loss in mistletoes. The increased water use of mistletoes resulted in significantly decreased host $\Psi _{\rm{leaf}}$ and $\Psi _{\rm{trunk}}$. It further translated to an estimated increase of up to 11% PLC for infected hosts, confirming greater hydraulic dysfunction of infected trees that place them at higher risk of hydraulic failure. However, uninfected branches of Eucalyptus fibrosa F.Muell. had much tighter controls on water loss than uninfected branches of Eucalyptus moluccana Roxb., which shifted the risk of hydraulic failure towards an increased risk of carbon starvation for E. fibrosa. The contrasting mechanistic responses to heat and drought stress between both co-occurring species demonstrates the complexity of host–parasite interactions and highlights the challenge in predicting species-specific responses to biotic agents in a warmer and drier climate.

Optimal stomatal theory predicts that stomata operate to maximise photosynthesis (A

Rising atmospheric [CO2 ] (Ca ) 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 Ca . Here, we sought to evaluate how increased vapor pressure deficit and reduced precipitation are likely to modify the impact of elevated Ca (eCa ) 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 Ca (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 Ca 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 Ca 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 Ca 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.

Despite knowledge of the interaction between climate change factors significant uncertainty exists concerning the individual and interactive effects of elevated carbon dioxide (eCO2) and elevated temperature (eT) on the soil microbiome and function. Here we examine the individual and interactive effects of eCO2 and eT on tree growth, soil respiration (Rsoil), biomass, structural and functional composition of microbial community, nitrogen (N) mineralisation and N availability in a whole tree chamber experiment. Eucalyptus globulus plants were grown from seedling to ca. 10 m tall for 15 months in a nutrient-poor sandy soil under ambient and elevated (+ 240 ppm) atmospheric CO2 concentrations combined with ambient or elevated temperatures (+ 3 °C) in a full factorial design. Plant growth was strongly stimulated under eCO2, but eT had little impact on any measured plant property. In contrast, Rsoil was not consistently affected by eCO2 or eT, but correlated strongly with root and leaf biomass. The response of N-mineralisation and nutrient availability to eCO2 and eT varied across time, and available N correlated strongly with plant height. Further, the C:N ratio of the microbial biomass and leaves were both higher under eCeT treatment. However, these functional measures were not significantly linked to either structural or functional diversity of the soil microbiome. Taken together, these results suggest that in this low-nutrient soil, belowground processes are principally driven by aboveground productivity. Our work provides novel insight into mechanisms underlying above- and belowground response to climate change, and the potential to sequester C in a low-nutrient status soil under future climatic conditions may be limited .

Ecosystems integrity and services are threatened by anthropogenic global changes. Mitigating and adapting to these changes require knowledge of ecosystem functioning in the expected novel environments, informed in large part through experimentation and modelling. This paper describes 13 advanced controlled environment facilities for experimental ecosystem studies, herein termed ecotrons, open to the international community. Ecotrons enable simulation of a wide range of natural environmental conditions in replicated and independent experimental units while measuring various ecosystem processes. This capacity to realistically control ecosystem environments is used to emulate a variety of climatic scenarios and soil conditions, in natural sunlight or through broad‐spectrum lighting. The use of large ecosystem samples, intact or reconstructed, minimizes border effects and increases biological and physical complexity. Measurements of concentrations of greenhouse trace gases as well as their net exchange between the ecosystem and the atmosphere are performed in most ecotrons, often quasi continuously. The flow of matter is often tracked with the use of stable isotope tracers of carbon and other elements. Equipment is available for measurements of soil water status as well as root and canopy growth. The experiments ran so far emphasize the diversity of the hosted research. Half of them concern global changes, often with a manipulation of more than one driver. About a quarter deal with the impact of biodiversity loss on ecosystem functioning and one quarter with ecosystem or plant physiology. We discuss how the methodology for environmental simulation and process measurements, especially in soil, can be improved and stress the need to establish stronger links with modelling in future projects. These developments will enable further improvements in mechanistic understanding and predictive capacity of ecotron research which will play, in complementarity with field experimentation and monitoring, a crucial role in exploring the ecosystem consequences of environmental changes., Experimentation and modelling are necessary to predict ecosystem functioning under future environments and to develop mitigating and adapting strategies. This paper describes 13 advanced controlled environment facilities, called ecotrons, open to the international community. An ecotron comprises a set of replicated enclosures designed to host ecosystems samples and enable realistic simulations of above‐ and belowground environmental conditions, while simultaneously and automatically measuring ecosystem processes. The characteristics of these infrastructures are given as well as examples of collaborative projects hosted so far.

Determining the relationship between reductions in stomatal conductance (g

Thermoregulation of leaf temperature (Tleaf ) may foster metabolic homeostasis in plants, but the degree to which Tleaf is moderated, and under what environmental contexts, is a topic of debate. Isotopic studies inferred the temperature of photosynthetic carbon assimilation to be a constant value of c. 20°C; by contrast, leaf biophysical theory suggests a strong dependence of Tleaf on environmental drivers. Can this apparent disparity be reconciled? We continuously measured Tleaf and whole-crown net CO2 uptake for Eucalyptus parramattensis trees growing in field conditions in whole-tree chambers under ambient and +3°C warming conditions, and calculated assimilation-weighted leaf temperature (TL-AW ) across 265 d, varying in air temperature (Tair ) from -1 to 45°C. We compared these data to TL-AW derived from wood cellulose δ18 O. Tleaf exhibited substantial variation driven by Tair , light intensity, and vapor pressure deficit, and Tleaf was strongly linearly correlated with Tair with a slope of c. 1.0. TL-AW values calculated from cellulose δ18 O vs crown fluxes were remarkably consistent; both varied seasonally and in response to the warming treatment, tracking variation in Tair . The leaves studied here were nearly poikilothermic, with no evidence of thermoregulation of Tleaf towards a homeostatic value. Importantly, this work supports the use of cellulose δ18 O to infer TL-AW , but does not support the concept of strong homeothermic regulation of Tleaf.

Plant respiration can acclimate to changing environmental conditions and vary between species as well as biome types, although belowground respiration responses to ongoing climate warming are not well understood. Understanding the thermal acclimation capacity of root respiration (Rroot) in relation to increasing temperatures is therefore critical in elucidating a key uncertainty in plant function in response to warming. However, the degree of temperature acclimation of Rroot in rainforest trees and how root chemical and morphological traits are related to acclimation is unknown. Here we investigated the extent to which respiration of fine roots (≤2 mm) of four tropical and four warm-temperate rainforest tree seedlings differed in response to warmer growth temperatures (control and +6 °C), including temperature sensitivity (Q10) and the degree of acclimation of Rroot. Regardless of biome type, we found no consistent pattern in the short-term temperature responses of Rroot to elevated growth temperature: a significant reduction in the temperature response of Rroot to +6 °C treatment was only observed for a tropical species, Cryptocarya mackinnoniana, whereas the other seven species had either some stimulation or no alteration. Across species, Rroot was positively correlated with root tissue nitrogen concentration (mg g−1), while Q10 was positively correlated with root tissue density (g cm−3). Warming increased root tissue density by 20.8% but did not alter root nitrogen across species. We conclude that thermal acclimation capacity of Rroot to warming is species-specific and suggest that root tissue density is a useful predictor of Rroot and its thermal responses in rainforest tree seedlings.

Shifts in the timing, intensity and/or frequency of climate extremes, such as severe drought and heatwaves, can generate sustained shifts in ecosystem function with important ecological and economic impacts for rangelands and managed pastures. The Pastures and Climate Extremes experiment (PACE) in Southeast Australia was designed to investigate the impacts of a severe winter/spring drought (60% rainfall reduction) and, for a subset of species, a factorial combination of drought and elevated temperature (ambient +3°C) on pasture productivity. The experiment included nine common pasture and Australian rangeland species from three plant functional groups (C3 grasses, C4 grasses and legumes) planted in monoculture. Winter/spring drought resulted in productivity declines of 45% on average and up to 74% for the most affected species (Digitaria eriantha) during the 6-month treatment period, with eight of the nine species exhibiting significant yield reductions. Despite considerable variation in species’ sensitivity to drought, C4 grasses were more strongly affected by this treatment than C3 grasses or legumes. Warming also had negative effects on cool-season productivity, associated at least partially with exceedance of optimum growth temperatures in spring and indirect effects on soil water content. The combination of winter/spring drought and year-round warming resulted in the greatest yield reductions. We identified responses that were either additive (Festuca), or less-than-additive (Medicago), where warming reduced the magnitude of drought effects. Results from this study highlight the sensitivity of diverse pasture species to increases in winter and spring drought severity similar to those predicted for this region, and that anticipated benefits of cool-season warming are unlikely to be realized. Overall, the substantial negative impacts on productivity suggest that future, warmer, drier climates will result in shortfalls in cool-season forage availability, with profound implications for the livestock industry and natural grazer communities.

Autotrophic respiration is a major driver of the global C cycle and may contribute a positive climate warming feedback through increased atmospheric concentrations of CO2 . The extent of this feedback depends on plants' ability to acclimate respiration to maintain a constant carbon use efficiency (CUE). We quantified respiratory partitioning of gross primary production (GPP) and CUE of field-grown trees in a long-term warming experiment (+3°C). We delivered a 13 C-CO2 pulse to whole tree crowns and chased that pulse in the respiration of leaves, whole crowns, roots, and soil. We also measured the isotopic composition of soil microbial biomass and the respiration rates of leaves and whole crowns. We documented homeostatic respiratory acclimation of foliar and whole-crown respiration rates; the trees adjusted to experimental warming such that leaf-level respiration rates were not increased. Experimental warming had no detectable impact on respiratory partitioning or mean residence times. Of the 13 C label acquired by the trees, aboveground respiration consumed 10%, belowground respiration consumed 40%, and the remaining 50% was retained. Experimental warming of +3°C did not alter respiratory partitioning at the scale of entire trees, suggesting that complete acclimation of respiration to warming is likely to dampen a positive climate warming feedback.

The allocation of carbon (C) is an important component of tree physiology that influences growth and ecosystem C storage. Allocation is challenging to measure, and its sensitivity to environmental changes such as warming and altered water availability is uncertain. We exposed young Eucalyptus tereticornis trees to +3°C warming and elimination of summer precipitation in the field using whole-tree chambers. We calculated C allocation terms using detailed measurements of growth and continuous whole-crown CO2 and water exchange measurements. Trees grew from small saplings to nearly 9 m height during this 15-month experiment. Warming accelerated growth and leaf area development, and it increased the partitioning of gross primary production (GPP) to aboveground respiration and growth while decreasing partitioning below ground. Eliminating summer precipitation reduced C gain and growth but did not impact GPP partitioning. Trees utilized deep soil water and avoided strongly negative water potentials. Warming increased growth respiration, but maintenance respiration acclimated homeostatically. The increasing growth in the warmed treatment resulted in higher rates of respiration, even with complete acclimation of maintenance respiration. Warming-induced stimulations of tree growth likely involve increased C allocation above ground, particularly to leaf area development, whereas reduced water availability may not stimulate allocation to roots.

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 ��mol 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 ��mol 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 Ca 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.

Shifts in the timing, intensity and/or frequency of climate extremes, such as severe drought and heatwaves, can generate sustained shifts in ecosystem function with important ecological and economic impacts for rangelands and managed pastures. The Pastures and Climate Extremes experiment (PACE) in Southeast Australia was designed to investigate the impacts of a severe winter/spring drought (60% rainfall reduction) and, for a subset of species, a factorial combination of drought and elevated temperature (ambient +3 °C) on pasture productivity. The experiment included nine common pasture and Australian rangeland species from three plant functional groups (C3 grasses, C4 grasses and legumes) planted in monoculture. Winter/spring drought resulted in productivity declines of 45% on average and up to 74% for the most affected species (Digitaria eriantha) during the 6-month treatment period, with eight of the nine species exhibiting significant yield reductions. Despite considerable variation in species’ sensitivity to drought, C4 grasses were more strongly affected by this treatment than C3 grasses or legumes. Warming also had negative effects on cool-season productivity, associated at least partially with exceedance of optimum growth temperatures in spring and indirect effects on soil water content. The combination of winter/spring drought and year-round warming resulted in the greatest yield reductions. We identified responses that were either additive such that there was only as significant warming effect under drought (Festuca), or less-than-additive, where there was no drought effect under warming (Medicago), compared to ambient plots. Results from this study highlight the sensitivity of diverse pasture species to increases in winter and spring drought severity similar to those predicted for this region, and that anticipated benefits of cool-season warming are unlikely to be realised. Overall, the substantial negative impacts on productivity suggest that future, warmer, drier climates will result in shortfalls in cool-season forage availability, with profound implications for the livestock industry and natural grazer communities.

Rising atmospheric [CO

Photosynthetic water-use efficiency (WUE) describes the link between terrestrial carbon (C) and water cycles. Estimates of intrinsic WUE (iWUE) from gas exchange and C isotopic composition (δ13 C) differ due to an internal conductance in the leaf mesophyll (gm ) that is variable and seldom computed. We present the first direct estimates of whole-tree gm , together with iWUE from whole-tree gas exchange and δ13 C of the phloem (δ13 Cph ). We measured gas exchange, online 13 C-discrimination, and δ13 Cph monthly throughout spring, summer, and autumn in Eucalyptus tereticornis grown in large whole-tree chambers. Six trees were grown at ambient temperatures and six at a 3°C warmer air temperature; a late-summer drought was also imposed. Drought reduced whole-tree gm . Warming had few direct effects, but amplified drought-induced reductions in whole-tree gm . Whole-tree gm was similar to leaf gm for these same trees. iWUE estimates from δ13 Cph agreed with iWUE from gas exchange, but only after incorporating gm . δ13 Cph was also correlated with whole-tree 13 C-discrimination, but offset by -2.5 ± 0.7‰, presumably due to post-photosynthetic fractionations. We conclude that δ13 Cph is a good proxy for whole-tree iWUE, with the caveats that post-photosynthetic fractionations and intrinsic variability of gm should be incorporated to provide reliable estimates of this trait in response to abiotic stress.

Xylem sap velocity of two dominant tree species, Eucalyptus crebra F. Muell. and Callitris glaucophylla J. Thompson & L.A.S. Johnson, in a native remnant forest of eastern Australia was measured in winter and summer during a prolonged (> 12 months) and extensive drought. The influence of vapour pressure deficit (VPD) and solar radiation levels on the velocity of sap was determined. Pronounced hysteresis in sap velocity was observed in both species as a function of VPD and solar radiation. However, the rotation of the hysteresis curve was clockwise for the response of sap velocity to VPD but anti-clockwise in the response of sap velocity to radiation levels. A possible reason for this difference is discussed.The degree of hysteresis (area bounded by the curve) was larger for the VPD response than the response to solar radiation and also varied with season. A simple linear model was able to predict sap velocity from knowledge of VPD and solar radiation in winter and summer. The consistent presence of hysteresis in the response to sap velocity to VPD and solar radiation suggests that large temporal and spatial models of vegetation water use may require some provision for the different responses of sap velocity, and hence water use, to VPD and solar radiation, between morning and afternoon and between seasons.

Atmospheric carbon dioxide enrichment (eCO2) can enhance plant carbon uptake and growth1–5, thereby providing an important negative feedback to climate change by slowing the rate of increase of the atmospheric CO2 concentration6. Although evidence gathered from young aggrading forests has generally indicated a strong CO2 fertilization effect on biomass growth3–5, it is unclear whether mature forests respond to eCO2 in a similar way. In mature trees and forest stands7–10, photosynthetic uptake has been found to increase under eCO2 without any apparent accompanying growth response, leaving the fate of additional carbon fixed under eCO2 unclear4,5,7–11. Here using data from the first ecosystem-scale Free-Air CO2 Enrichment (FACE) experiment in a mature forest, we constructed a comprehensive ecosystem carbon budget to track the fate of carbon as the forest responded to four years of eCO2 exposure. We show that, although the eCO2 treatment of +150 parts per million (+38 per cent) above ambient levels induced a 12 per cent (+247 grams of carbon per square metre per year) increase in carbon uptake through gross primary production, this additional carbon uptake did not lead to increased carbon sequestration at the ecosystem level. Instead, the majority of the extra carbon was emitted back into the atmosphere via several respiratory fluxes, with increased soil respiration alone accounting for half of the total uptake surplus. Our results call into question the predominant thinking that the capacity of forests to act as carbon sinks will be generally enhanced under eCO2, and challenge the efficacy of climate mitigation strategies that rely on ubiquitous CO2 fertilization as a driver of increased carbon sinks in global forests. Carbon dioxide enrichment of a mature forest resulted in the emission of the excess carbon back into the atmosphere via enhanced ecosystem respiration, suggesting that mature forests may be limited in their capacity to mitigate climate change.

Thermoregulation of leaf temperature (T

Predicting the seasonal dynamics of ecosystem carbon fluxes is challenging in broadleaved evergreen forests because of their moderate climates and subtle changes in canopy phenology. We assessed the climatic and biotic drivers of the seasonality of net ecosystem–atmosphere CO2 exchange (NEE) of a eucalyptus-dominated forest near Sydney, Australia, using the eddy covariance method. The climate is characterised by a mean annual precipitation of 800 mm and a mean annual temperature of 18 ∘C, hot summers and mild winters, with highly variable precipitation. In the 4-year study, the ecosystem was a sink each year (−225 g C m−2 yr−1 on average, with a standard deviation of 108 g C m−2 yr−1); inter-annual variations were not related to meteorological conditions. Daily net C uptake was always detected during the cooler, drier winter months (June through August), while net C loss occurred during the warmer, wetter summer months (December through February). Gross primary productivity (GPP) seasonality was low, despite longer days with higher light intensity in summer, because vapour pressure deficit (D) and air temperature (Ta) restricted surface conductance during summer while winter temperatures were still high enough to support photosynthesis. Maximum GPP during ideal environmental conditions was significantly correlated with remotely sensed enhanced vegetation index (EVI; r2 = 0.46) and with canopy leaf area index (LAI; r2 = 0.29), which increased rapidly after mid-summer rainfall events. Ecosystem respiration (ER) was highest during summer in wet soils and lowest during winter months. ER had larger seasonal amplitude compared to GPP, and therefore drove the seasonal variation of NEE. Because summer carbon uptake may become increasingly limited by atmospheric demand and high temperature, and because ecosystem respiration could be enhanced by rising temperatures, our results suggest the potential for large-scale seasonal shifts in NEE in sclerophyll vegetation under climate change.

Predicting the seasonal dynamics of ecosystem carbon fluxes is challenging in broadleaved evergreen forests because of their moderate climates and subtle changes in canopy phenology. We assessed the climatic and biotic drivers of the seasonality of net ecosystem–atmosphere CO2 exchange (NEE) of a eucalyptus-dominated forest near Sydney, Australia, using the eddy covariance method. The climate is characterised by a mean annual precipitation of 800 mm and a mean annual temperature of 18 ∘C, hot summers and mild winters, with highly variable precipitation. In the 4-year study, the ecosystem was a sink each year (−225 g C m−2 yr−1 on average, with a standard deviation of 108 g C m−2 yr−1); inter-annual variations were not related to meteorological conditions. Daily net C uptake was always detected during the cooler, drier winter months (June through August), while net C loss occurred during the warmer, wetter summer months (December through February). Gross primary productivity (GPP) seasonality was low, despite longer days with higher light intensity in summer, because vapour pressure deficit (D) and air temperature (Ta) restricted surface conductance during summer while winter temperatures were still high enough to support photosynthesis. Maximum GPP during ideal environmental conditions was significantly correlated with remotely sensed enhanced vegetation index (EVI; r2 = 0.46) and with canopy leaf area index (LAI; r2 = 0.29), which increased rapidly after mid-summer rainfall events. Ecosystem respiration (ER) was highest during summer in wet soils and lowest during winter months. ER had larger seasonal amplitude compared to GPP, and therefore drove the seasonal variation of NEE. Because summer carbon uptake may become increasingly limited by atmospheric demand and high temperature, and because ecosystem respiration could be enhanced by rising temperatures, our results suggest the potential for large-scale seasonal shifts in NEE in sclerophyll vegetation under climate change.

The CO2 efflux from tree stem surfaces to atmosphere (RS) is an important component in the carbon (C) balance of forest ecosystems. Rising atmospheric carbon dioxide concentrations [CO2] are expected to stimulate RS, because of greater C assimilation and carbohydrate supply to stems under rising [CO2]. Growth respiration (Rg) and maintenance respiration (Rm) during the warm growing season may respond differently to rising [CO2] due to different metabolic demands. To test the effect of elevated [CO2] (eCO2, ambient +150 ppm) on RS, we examined RS in mature Eucalyptus trees on a monthly basis for an entire year during the seventh year of exposure to eCO2. RS varied seasonally and mirrored seasonal variation in temperature. While RS was not significantly increased under eCO2 compared to ambient CO2 (aCO2), its temperature sensitivity was significantly decreased (Q10 of 1.92 for aCO2 and 1.56 for eCO2). The estimated annual Rg accounted for approximately 7–8% of annual total RS, 419 ± 103 g C m−2 yr-1, indicating that Rm contributes substantially to total RS in this mature woodland. Monthly mean RS was correlated with monthly mean soil temperature, soil moisture and monthly stem growth rate in this dry year, but soil moisture levels may have been insufficient to observe the impacts of eCO2 on stem growth in this droughted and phosphorous limited site. Our results highlight that eCO2 tends to increase Rm at low temperatures during the non-growing season, thus decreasing the temperature sensitivity of RS, despite a neutral effect of eCO2 on RS rates on a yearly basis.