Urban greening with shrubs can supercharge invertebrate abundance and diversity.

In urban areas, diverse and complex habitats for biodiversity are often lacking. This lack of diversity not only compromises essential ecological processes, such as pollination and nutrient cycling, but also diminishes the resilience of urban ecosystems to pests and diseases. To enhance urban biodiversity, a possible solution is to integrate shrubs alongside trees, thereby increasing the overall amount of vegetation, structural complexity and the associated resource diversity. Here, using a common garden experiment involving a variety of trees and shrubs planted alone and in combination, we evaluate how canopy-associated invertebrate assemblages are influenced by vegetation type. In particular, we test whether the presence of shrubs, alone or with trees, results in increased abundance and taxonomic richness of invertebrates, compared to trees on their own. We found that the overall abundance of invertebrates, and that of specific functional groups (e.g., herbivores, pollinators, detritivores), was higher on shrubs, compared to trees, and when trees and shrubs were planted in combination (relative to trees on their own). Our results suggest that planting shrub and tree species with wide and dense crowns can increase the associated abundance and taxonomic and functional group richness of invertebrate communities. Overall, our findings indicate that urban planning would benefit from incorporating shrubs alongside urban trees to maximise invertebrate abundance, diversity and function in urban landscapes.

Decoupling between stomatal conductance and photosynthesis occurs under extreme heat in broadleaf tree species regardless of water access.

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.

Aridity drives clinal patterns in leaf traits and responsiveness to precipitation in a broadly distributed Australian tree species.

Aridity shapes species distributions and plant growth and function worldwide. Yet, plant traits often show complex relationships with aridity, challenging our understanding of aridity as a driver of evolutionary adaptation. We grew nine genotypes of Eucalyptus camaldulensis subsp. camaldulensis sourced from an aridity gradient together in the field for ~650 days under low and high precipitation treatments. Eucalyptus camaldulesis is considered a phreatophyte (deep-rooted species that utilizes groundwater), so we hypothesized that genotypes from more arid environments would show lower aboveground productivity, higher leaf gas-exchange rates, and greater tolerance/avoidance of dry surface soils (indicated by lower responsiveness) than genotypes from less arid environments. Aridity predicted genotype responses to precipitation, with more arid genotypes showing lower responsiveness to reduced precipitation and dry surface conditions than less arid genotypes. Under low precipitation, genotype net photosynthesis and stomatal conductance increased with home-climate aridity. Across treatments, genotype intrinsic water-use efficiency and osmotic potential declined with increasing aridity while photosynthetic capacity (Rubisco carboxylation and RuBP regeneration) increased with aridity. The observed clinal patterns indicate that E. camaldulensis genotypes from extremely arid environments possess a unique strategy defined by lower responsiveness to dry surface soils, low water-use efficiency, and high photosynthetic capacity. This strategy could be underpinned by deep rooting and could be adaptive under arid conditions where heat avoidance is critical and water demand is high.

Thermal acclimation of leaf respiration is consistent in tropical and subtropical populations of two mangrove species.

Populations from different climates often show unique growth responses to temperature, reflecting temperature adaptation. Yet, whether populations from different climates differ in physiological temperature acclimation remains unclear. Here, we test whether populations from differing thermal environments exhibit different growth responses to temperature and differences in temperature acclimation of leaf respiration. We grew tropical and subtropical populations of two mangrove species (Avicennia germinans and Rhizophora mangle) under ambient and experimentally warmed conditions in a common garden at the species' northern range limit. We quantified growth and temperature responses of leaf respiration (R) at seven time points over ~10 months. Warming increased productivity of tropical populations more than subtropical populations, reflecting a higher temperature optimum for growth. In both species, R measured at 25 °C declined as seasonal temperatures increased, demonstrating thermal acclimation. Contrary to our expectations, acclimation of R was consistent across populations and temperature treatments. However, populations differed in adjusting the temperature sensitivity of R (Q10) to seasonal temperatures. Following a freeze event, tropical Avicennia showed greater freeze damage than subtropical Avicennia, while both Rhizophora populations appeared equally susceptible. We found evidence of temperature adaptation at the whole-plant scale but little evidence for population differences in thermal acclimation of leaf physiology. Studies that examine potential costs and benefits of thermal acclimation in an evolutionary context may provide new insights into limits of thermal acclimation.

Belowground carbon allocation, root trait plasticity, and productivity during drought and warming in a pasture grass.

Sustaining grassland production in a changing climate requires an understanding of plant adaptation strategies, including trait plasticity under warmer and drier conditions. However, our knowledge to date disproportionately relies on aboveground responses, despite the importance of belowground traits in maintaining aboveground growth, especially in grazed systems. We subjected a perennial pasture grass, Festuca arundinacea, to year-round warming (+3 °C) and cool-season drought (60% rainfall reduction) in a factorial field experiment to test the hypotheses that: (i) drought and warming increase carbon allocation belowground and shift root traits towards greater resource acquisition and (ii) increased belowground carbon reserves support post-drought aboveground recovery. Drought and warming reduced plant production and biomass allocation belowground. Drought increased specific root length and reduced root diameter in warmed plots but increased root starch concentrations under ambient temperature. Higher diameter and soluble sugar concentrations of roots and starch storage in crowns explained aboveground production under climate extremes. However, the lack of association between post-drought aboveground biomass and belowground carbon and nitrogen reserves contrasted with our predictions. These findings demonstrate that root trait plasticity and belowground carbon reserves play a key role in aboveground production during climate stress, helping predict pasture responses and inform management decisions under future climates.

Nocturnal plant respiration is under strong non-temperature control.

Most biological rates depend on the rate of respiration. Temperature variation is typically considered the main driver of daily plant respiration rates, assuming a constant daily respiration rate at a set temperature. Here, we show empirical data from 31 species from temperate and tropical biomes to demonstrate that the rate of plant respiration at a constant temperature decreases monotonically with time through the night, on average by 25% after 8 h of darkness. Temperature controls less than half of the total nocturnal variation in respiration. A new universal formulation is developed to model and understand nocturnal plant respiration, combining the nocturnal decrease in the rate of plant respiration at constant temperature with the decrease in plant respiration according to the temperature sensitivity. Application of the new formulation shows a global reduction of 4.5 -6 % in plant respiration and an increase of 7-10% in net primary production for the present-day.

Crown dieback and mortality of urban trees linked to heatwaves during extreme drought.

Cities have been described as 'heat islands' and 'dry islands' due to hotter, drier air in urban areas, relative to the surrounding landscape. As climate change intensifies, the health of urban trees will be increasingly impacted. Here, we posed the question: Is it possible to predict urban tree species mortality using (1) species climate envelopes and (2) plant functional traits? To answer these, we tracked patterns of crown dieback and recovery for 23 common urban tree and shrub species in Sydney, Australia during the record-breaking austral 2019-2020 summer. We identified 10 heat-tolerant species including five native and five exotic species, which represent climate-resilient options for urban plantings that are likely to continue to thrive for decades. Thirteen species were considered vulnerable to adverse conditions due to their mortality, poor health leading to tree removal, and/or extensive crown dieback. Crown dieback increased with increasing precipitation of the driest month of species climate of origin, suggesting that species from dry climates may be better suited for urban forests in future climates. We effectively grouped species according to their drought strategy (i.e., tolerance versus avoidance) using a simple trait-based framework that was directly linked with species mortality. The seven most climate-vulnerable species used a drought-avoidance strategy, having low wood density and high turgor loss points along with large, thin leaves with low heat tolerance. Overall, plant functional traits were better than species climate envelopes at explaining crown dieback. Recovery after stress required two mild, wet years for most species, resulting in prolonged loss of cooling benefits as well as economic losses due to replacement of dead/damaged trees. Hotter, longer, and more frequent heatwaves will require selection of more climate-resilient species in urban forests, and our results suggest that future research should focus on plant thermal traits to improve prediction models and species selection.

Consistent diurnal pattern of leaf respiration in the light among contrasting species and climates.

Leaf daytime respiration (leaf respiration in the light, RL ) is often assumed to constitute a fixed fraction of leaf dark respiration (RD ) (i.e. a fixed light inhibition of respiration (RD )) and vary diurnally due to temperature fluctuations. These assumptions were tested by measuring RL , RD and the light inhibition of RD in the field at a constant temperature using the Kok method. Measurements were conducted diurnally on 21 different species: 13 deciduous, four evergreen and four herbaceous from humid continental and humid subtropical climates. RL and RD showed significant diurnal variations and the diurnal pattern differed in trajectory and magnitude between climates, but not between plant functional types (PFTs). The light inhibition of RD varied diurnally and differed between climates and in trajectory between PFTs. The results highlight the entrainment of leaf daytime respiration to the diurnal cycle and that time of day should be accounted for in studies seeking to examine the environmental and biological drivers of leaf daytime respiration.

Drought and warming alter gross primary production allocation and reduce productivity in a widespread pasture grass.

Carbon allocation determines plant growth, fitness and reproductive success. However, climate warming and drought impacts on carbon allocation patterns in grasses are not well known, particularly following grazing or clipping. A widespread C3 pasture grass, Festuca arundinacea, was grown at 26 and 30°C in controlled environment chambers and subjected to drought (65% reduction relative to well-watered controls). Leaf, root and whole-plant carbon fluxes were measured and linked to growth before and after clipping. Both drought and warming reduced gross primary production and plant biomass. Drought reduced net leaf photosynthesis but increased the leaf respiratory fraction of assimilated carbon. Warming increased root respiration but did not affect either net leaf photosynthesis or leaf respiration. There was no evidence of thermal acclimation. Moreover, root respiratory carbon loss was amplified in the combined drought and warming treatment and, in addition to a negative carbon balance aboveground, explained an enhanced reduction in plant biomass. Plant regrowth following clipping was strongly suppressed by drought, reflecting increased tiller mortality and exacerbated respiratory carbon loss. These findings emphasize the importance of considering carbon allocation patterns in response to grazing or clipping and interactions with climatic factors for sustainable pasture production in a future climate.

Pastures and Climate Extremes: Impacts of Cool Season Warming and Drought on the Productivity of Key Pasture Species in a Field Experiment.

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.

Tropical rainforest species have larger increases in temperature optima with warming than warm-temperate rainforest trees.

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 (ToptA ) than temperate species. There was acclimation of ToptA 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 Vcmax25 and Jmax25 . The temperature sensitivity of respiration (Q10 ) 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.

Extreme heat increases stomatal conductance and drought-induced mortality risk in vulnerable plant species.

Tree mortality during global-change-type drought is usually attributed to xylem dysfunction, but as climate change increases the frequency of extreme heat events, it is necessary to better understand the interactive role of heat stress. We hypothesized that some drought-stressed plants paradoxically open stomata in heatwaves to prevent leaves from critically overheating. We experimentally imposed heat (>40°C) and drought stress onto 20 broadleaf evergreen tree/shrub species in a glasshouse study. Most well-watered plants avoided lethal overheating, but drought exacerbated thermal damage during heatwaves. Thermal safety margins (TSM) quantifying the difference between leaf surface temperature and leaf critical temperature, where photosynthesis is disrupted, identified species vulnerability to heatwaves. Several mechanisms contributed to high heat tolerance and avoidance of damaging leaf temperatures-small leaf size, low leaf osmotic potential, high leaf mass per area (i.e., thick, dense leaves), high transpirational capacity, and access to water. Water-stressed plants had smaller TSM, greater crown dieback, and a fundamentally different stomatal heatwave response relative to well-watered plants. On average, well-watered plants closed stomata and decreased stomatal conductance (gs ) during the heatwave, but droughted plants did not. Plant species with low gs , either due to isohydric stomatal behavior under water deficit or inherently low transpirational capacity, opened stomata and increased gs under high temperatures. The current paradigm maintains that stomata close before hydraulic thresholds are surpassed, but our results suggest that isohydric species may dramatically increase gs (over sixfold increases) even past their leaf turgor loss point. By actively increasing water loss at high temperatures, plants can be driven toward mortality thresholds more rapidly than has been previously recognized. The inclusion of TSM and responses to heat stress could improve our ability to predict the vulnerability of different tree species to future droughts.

Increasing aridity will not offset CO2 fertilization in fast-growing eucalypts with access to deep soil water.

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.

Acclimation of leaf respiration temperature responses across thermally contrasting biomes.

Short-term temperature response curves of leaf dark respiration (R-T) provide insights into a critical process that influences plant net carbon exchange. This includes how respiratory traits acclimate to sustained changes in the environment. Our study analysed 860 high-resolution R-T (10-70°C range) curves for: (a) 62 evergreen species measured in two contrasting seasons across several field sites/biomes; and (b) 21 species (subset of those sampled in the field) grown in glasshouses at 20°C : 15°C, 25°C : 20°C and 30°C : 25°C, day : night. In the field, across all sites/seasons, variations in R25 (measured at 25°C) and the leaf T where R reached its maximum (Tmax ) were explained by growth T (mean air-T of 30-d before measurement), solar irradiance and vapour pressure deficit, with growth T having the strongest influence. R25 decreased and Tmax increased with rising growth T across all sites and seasons with the single exception of winter at the cool-temperate rainforest site where irradiance was low. The glasshouse study confirmed that R25 and Tmax thermally acclimated. Collectively, the results suggest: (1) thermal acclimation of leaf R is common in most biomes; and (2) the high T threshold of respiration dynamically adjusts upward when plants are challenged with warmer and hotter climates.

Whole-tree mesophyll conductance reconciles isotopic and gas-exchange estimates of water-use efficiency.

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.

Ecotrons: Powerful and versatile ecosystem analysers for ecology, agronomy and environmental science.

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.

No evidence of homeostatic regulation of leaf temperature in Eucalyptus parramattensis trees: integration of CO2 flux and oxygen isotope methodologies.

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.

Does root respiration in Australian rainforest tree seedlings acclimate to experimental warming?

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.

The fate of carbon in a mature forest under carbon dioxide enrichment.

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.