Microbial competition for phosphorus limits the CO2 response of a mature forest.

The capacity for terrestrial ecosystems to sequester additional carbon (C) with rising CO2 concentrations depends on soil nutrient availability1,2. Previous evidence suggested that mature forests growing on phosphorus (P)-deprived soils had limited capacity to sequester extra biomass under elevated CO2 (refs. 3-6), but uncertainty about ecosystem P cycling and its CO2 response represents a crucial bottleneck for mechanistic prediction of the land C sink under climate change7. Here, by compiling the first comprehensive P budget for a P-limited mature forest exposed to elevated CO2, we show a high likelihood that P captured by soil microorganisms constrains ecosystem P recycling and availability for plant uptake. Trees used P efficiently, but microbial pre-emption of mineralized soil P seemed to limit the capacity of trees for increased P uptake and assimilation under elevated CO2 and, therefore, their capacity to sequester extra C. Plant strategies to stimulate microbial P cycling and plant P uptake, such as increasing rhizosphere C release to soil, will probably be necessary for P-limited forests to increase C capture into new biomass. Our results identify the key mechanisms by which P availability limits CO2 fertilization of tree growth and will guide the development of Earth system models to predict future long-term C storage.

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.

Pyrogeography in flux: Reorganization of Australian fire regimes in a hotter world.

Changes to the spatiotemporal patterns of wildfire are having profound implications for ecosystems and society globally, but we have limited understanding of the extent to which fire regimes will reorganize in a warming world. While predicting regime shifts remains challenging because of complex climate-vegetation-fire feedbacks, understanding the climate niches of fire regimes provides a simple way to identify locations most at risk of regime change. Using globally available satellite datasets, we constructed 14 metrics describing the spatiotemporal dimensions of fire and then delineated Australia's pyroregions-the geographic area encapsulating a broad fire regime. Cluster analysis revealed 18 pyroregions, notably including the (1) high-intensity, infrequent fires of the temperate forests, (2) high-frequency, smaller fires of the tropical savanna, and (3) low-intensity, diurnal, human-engineered fires of the agricultural zones. To inform the risk of regime shifts, we identified locations where the climate under three CMIP6 scenarios is projected to shift (i) beyond each pyroregion's historical climate niche, and (ii) into climate space that is novel to the Australian continent. Under middle-of-the-road climate projections (SSP2-4.5), an average of 65% of the extent of the pyroregions occurred beyond their historical climate niches by 2081-2100. Further, 52% of pyroregion extents, on average, were projected to occur in climate space without present-day analogues on the Australian continent, implying high risk of shifting to states that also lack present-day counterparts. Pyroregions in tropical and hot-arid climates were most at risk of shifting into both locally and continentally novel climate space because (i) their niches are narrower than southern temperate pyroregions, and (ii) their already-hot climates lead to earlier departure from present-day climate space. Such a shift implies widespread risk of regime shifts and the emergence of no-analogue fire regimes. Our approach can be applied to other regions to assess vulnerability to rapid fire regime change.

Hydraulic failure and tree size linked with canopy die-back in eucalypt forest during extreme drought.

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.

Hydraulic and photosynthetic limitations prevail over root non-structural carbohydrate reserves as drivers of resprouting in two Mediterranean oaks.

Resprouting is an ancestral trait in angiosperms that confers resilience after perturbations. As climate change increases stress, resprouting vigor is declining in many forest regions, but the underlying mechanism is poorly understood. Resprouting in woody plants is thought to be primarily limited by the availability of non-structural carbohydrate reserves (NSC), but hydraulic limitations could also be important. We conducted a multifactorial experiment with two levels of light (ambient, 2-3% of ambient) and three levels of water stress (0, 50 and 80 percent losses of hydraulic conductivity, PLC) on two Mediterranean oaks (Quercus ilex and Q. faginea) under a rain-out shelter (n = 360). The proportion of resprouting individuals after canopy clipping declined markedly as PLC increased for both species. NSC concentrations affected the response of Q. ilex, the species with higher leaf construction costs, and its effect depended on the PLC. The growth of resprouting individuals was largely dependent on photosynthetic rates for both species, while stored NSC availability and hydraulic limitations played minor and non-significant roles, respectively. Contrary to conventional wisdom, our results indicate that resprouting in oaks may be primarily driven by complex interactions between hydraulics and carbon sources, whereas stored NSC play a significant but secondary role.

The validity of optimal leaf traits modelled on environmental conditions.

The ratio of leaf intercellular to ambient CO2 (χ) is modulated by stomatal conductance (gs ). These quantities link carbon (C) assimilation with transpiration, and along with photosynthetic capacities (Vcmax and Jmax ) are required to model terrestrial C uptake. We use optimization criteria based on the growth environment to generate predicted values of photosynthetic and water-use efficiency traits and test these against a unique dataset. Leaf gas-exchange parameters and carbon isotope discrimination were analysed in relation to local climate across a continental network of study sites. Sun-exposed leaves of 50 species at seven sites were measured in contrasting seasons. Values of χ predicted from growth temperature and vapour pressure deficit were closely correlated to ratios derived from C isotope (δ13 C) measurements. Correlations were stronger in the growing season. Predicted values of photosynthetic traits, including carboxylation capacity (Vcmax ), derived from δ13 C, growth temperature and solar radiation, showed meaningful agreement with inferred values derived from gas-exchange measurements. Between-site differences in water-use efficiency were, however, only weakly linked to the plant's growth environment and did not show seasonal variation. These results support the general hypothesis that many key parameters required by Earth system models are adaptive and predictable from plants' growth environments.

Understorey productivity in temperate grassy woodland responds to soil water availability but not to elevated [CO2 ].

Rising atmospheric [CO2 ] and associated climate change are expected to modify primary productivity across a range of ecosystems globally. Increasing aridity is predicted to reduce grassland productivity, although rising [CO2 ] and associated increases in plant water use efficiency may partially offset the effect of drying on growth. Difficulties arise in predicting the direction and magnitude of future changes in ecosystem productivity, due to limited field experimentation investigating climate and CO2 interactions. We use repeat near-surface digital photography to quantify the effects of water availability and experimentally manipulated elevated [CO2 ] (eCO2 ) on understorey live foliage cover and biomass over three growing seasons in a temperate grassy woodland in south-eastern Australia. We hypothesised that (i) understorey herbaceous productivity is dependent upon soil water availability, and (ii) that eCO2 will increase productivity, with greatest stimulation occurring under conditions of low water availability. Soil volumetric water content (VWC) determined foliage cover and growth rates over the length of the growing season (August to March), with low VWC (<0.1 m3  m-3 ) reducing productivity. However, eCO2 did not increase herbaceous cover and biomass over the duration of the experiment, or mitigate the effects of low water availability on understorey growth rates and cover. Our findings suggest that projected increases in aridity in temperate woodlands are likely to lead to reduced understorey productivity, with little scope for eCO2 to offset these changes.

Canopy leaf area of a mature evergreen Eucalyptus woodland does not respond to elevated atmospheric [CO2] but tracks water availability.

Canopy leaf area, quantified by the leaf area index (L), is a crucial driver of forest productivity, water use and energy balance. Because L responds to environmental drivers, it can represent an important feedback to climate change, but its responses to rising atmospheric [CO2] and water availability of forests have been poorly quantified. We studied canopy leaf area dynamics for 28 months in a native evergreen Eucalyptus woodland exposed to free-air CO2 enrichment (the EucFACE experiment), in a subtropical climate where water limitation is common. We hypothesized that, because of expected stimulation of productivity and water-use efficiency, L should increase with elevated [CO2]. We estimated L from diffuse canopy transmittance, and measured monthly leaf litter production. Contrary to expectation, L did not respond to elevated [CO2]. We found that L varied between 1.10 and 2.20 across the study period. The dynamics of L showed a quick increase after heavy rainfall and a steady decrease during periods of low rainfall. Leaf litter production was correlated to changes in L, both during periods of decreasing L (when no leaf growth occurred) and during periods of increasing L (active shedding of old foliage when new leaf growth occurred). Leaf lifespan, estimated from mean L and total annual litter production, was up to 2 months longer under elevated [CO2] (1.18 vs. 1.01 years; P = 0.05). Our main finding that L was not responsive to elevated CO2 is consistent with other forest FACE studies, but contrasts with the positive response of L commonly predicted by many ecosystem models.

Environmental correlates of the forest carbon distribution in the Central Himalayas.

Understanding the biophysical limitations on forest carbon across diverse ecological regions is crucial for accurately assessing and managing forest carbon stocks. This study investigates the role of climate and disturbance on the spatial variation of two key forest carbon pools: aboveground carbon (AGC) and soil organic carbon (SOC). Using plot-level carbon pool estimates from Nepal's national forest inventory and structural equation modelling, we explore the relationship of forest carbon stocks to broad-scale climatic water and energy availability and fine-scale terrain and disturbance. The forest AGC and SOC models explained 25% and 59% of the observed spatial variation in forest AGC and SOC, respectively. Among the evaluated variables, disturbance exhibited the strongest negative correlation with AGC, while the availability of climatic energy demonstrated the strongest negative correlation with SOC. Disturbances such as selective logging and firewood collection result in immediate forest carbon loss, while soil carbon changes take longer to respond. The lower decomposition rates in the high-elevation region, due to lower temperatures, preserve organic matter and contribute to the high SOC stocks observed there. These results highlight the critical role of climate and disturbance regimes in shaping landscape patterns of forest carbon stocks. Understanding the underlying drivers of these patterns is crucial for forest carbon management and conservation across diverse ecological zones including the Central Himalayas.

A monthly gridded burned area database of national wildland fire data.

We assembled the first gridded burned area (BA) database of national wildfire data (ONFIRE), a comprehensive and integrated resource for researchers, non-government organisations, and government agencies analysing wildfires in various regions of the Earth. We extracted and harmonised records from different regions and sources using open and reproducible methods, providing data in a common framework for the whole period available (starting from 1950 in Australia, 1959 in Canada, 1985 in Chile, 1980 in Europe, and 1984 in the United States) up to 2021 on a common 1° × 1° grid. The data originate from national agencies (often, ground mapping), thus representing the best local expert knowledge. Key opportunities and limits in using this dataset are discussed as well as possible future expansions of this open-source approach that should be explored. This dataset complements existing gridded BA data based on remote sensing and offers a valuable opportunity to better understand and assess fire regime changes, and their drivers, in these regions. The ONFIRE database can be freely accessed at https://zenodo.org/record/8289245 .

Carbon-phosphorus cycle models overestimate CO2 enrichment response in a mature Eucalyptus forest.

The importance of phosphorus (P) in regulating ecosystem responses to climate change has fostered P-cycle implementation in land surface models, but their CO2 effects predictions have not been evaluated against measurements. Here, we perform a data-driven model evaluation where simulations of eight widely used P-enabled models were confronted with observations from a long-term free-air CO2 enrichment experiment in a mature, P-limited Eucalyptus forest. We show that most models predicted the correct sign and magnitude of the CO2 effect on ecosystem carbon (C) sequestration, but they generally overestimated the effects on plant C uptake and growth. We identify leaf-to-canopy scaling of photosynthesis, plant tissue stoichiometry, plant belowground C allocation, and the subsequent consequences for plant-microbial interaction as key areas in which models of ecosystem C-P interaction can be improved. Together, this data-model intercomparison reveals data-driven insights into the performance and functionality of P-enabled models and adds to the existing evidence that the global CO2-driven carbon sink is overestimated by models.

Woody clockworks: circadian regulation of night-time water use in Eucalyptus globulus.

The role of the circadian clock in controlling the metabolism of entire trees has seldom been considered. We tested whether the clock influences nocturnal whole-tree water use. Whole-tree chambers allowed the control of environmental variables (temperature, relative humidity). Night-time stomatal conductance (gs ) and sap flow (Q) were monitored in 6- to 8-m-tall Eucalyptus globulus trees during nights when environmental variables were kept constant, and also when conditions varied with time. Artificial neural networks were used to quantify the relative importance of circadian regulation of gs and Q. Under a constant environment, gs and Q declined from 0 to 6 h after dusk, but increased from 6 to 12 h after dusk. While the initial decline could be attributed to multiple processes, the subsequent increase is most consistent with circadian regulation of gs and Q. We conclude that endogenous regulation of gs is an important driver of night-time Q under natural environmental variability. The proportion of nocturnal Q variation associated with circadian regulation (23-56%) was comparable to that attributed to vapor pressure deficit variation (25-58%). This study contributes to our understanding of the linkages between molecular and cellular processes related to circadian regulation, and whole-tree processes related to ecosystem gas exchange in the field.

Plot-level estimates of aboveground biomass and soil organic carbon stocks from Nepal's forest inventory.

Given the contribution of deforestation and forest degradation to the global carbon cycle, forest resources are critical to mitigating the global climate change effects. Improved forest monitoring across different biomes is important to understand forest dynamics better and improve global projections of future atmospheric CO2 concentration. Better quantification of the forest carbon cycle advances scientific understanding and informs global negotiations about carbon emissions reduction. High-quality estimates of forest carbon stocks are currently scarce in many developing countries. Here, we present the most comprehensive georeferenced data set to date of plot-level forest carbon estimates for Nepal. Based on field observations from Nepal's national forest inventory of 2010-2014; the data set includes estimates for two major forest carbon pools, aboveground biomass (AGB) and soil organic carbon (SOC) stocks from 2,009 and 1,156 inventory plots, respectively. The dataset fills an important knowledge gap about forest carbon stocks in the Central Himalayas, a region with highly heterogeneous environmental conditions and rich biodiversity that is poorly represented in existing global estimates of forest carbon.

Mapping soil organic carbon stocks in Nepal's forests.

Comprehensive forest carbon accounting requires reliable estimation of soil organic carbon (SOC) stocks. Despite being an important carbon pool, limited information is available on SOC stocks in global forests, particularly for forests in mountainous regions, such as the Central Himalayas. The availability of consistently measured new field data enabled us to accurately estimate forest soil organic carbon (SOC) stocks in Nepal, addressing a previously existing knowledge gap. Our method involved modelling plot-based estimates of forest SOC using covariates related to climate, soil, and topographic position. Our quantile random forest model resulted in the high spatial resolution prediction of Nepal's national forest SOC stock together with prediction uncertainties. Our spatially explicit forest SOC map showed the high SOC levels in high-elevation forests and a significant underrepresentation of these stocks in global-scale assessments. Our results offer an improved baseline on the distribution of total carbon in the forests of the Central Himalayas. The benchmark maps of predicted forest SOC and associated errors, along with our estimate of 494 million tonnes (SE = 16) of total SOC in the topsoil (0-30 cm) of forested areas in Nepal, carry important implications for understanding the spatial variability of forest SOC in mountainous regions with complex terrains.

The 2019-2020 Australian forest fires are a harbinger of decreased prescribed burning effectiveness under rising extreme conditions.

There is an imperative for fire agencies to quantify the potential for prescribed burning to mitigate risk to life, property and environmental values while facing changing climates. The 2019-2020 Black Summer fires in eastern Australia raised questions about the effectiveness of prescribed burning in mitigating risk under unprecedented fire conditions. We performed a simulation experiment to test the effects of different rates of prescribed burning treatment on risks posed by wildfire to life, property and infrastructure. In four forested case study landscapes, we found that the risks posed by wildfire were substantially higher under the fire weather conditions of the 2019-2020 season, compared to the full range of long-term historic weather conditions. For area burnt and house loss, the 2019-2020 conditions resulted in more than a doubling of residual risk across the four landscapes, regardless of treatment rate (mean increase of 230%, range 164-360%). Fire managers must prepare for a higher level of residual risk as climate change increases the likelihood of similar or even more dangerous fire seasons.

Canopy dieback and recovery in Australian native forests following extreme drought.

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.

Forest fire threatens global carbon sinks and population centres under rising atmospheric water demand.

Levels of fire activity and severity that are unprecedented in the instrumental record have recently been observed in forested regions around the world. Using a large sample of daily fire events and hourly climate data, here we show that fire activity in all global forest biomes responds strongly and predictably to exceedance of thresholds in atmospheric water demand, as measured by maximum daily vapour pressure deficit. The climatology of vapour pressure deficit can therefore be reliably used to predict forest fire risk under projected future climates. We find that climate change is projected to lead to widespread increases in risk, with at least 30 additional days above critical thresholds for fire activity in forest biomes on every continent by 2100 under rising emissions scenarios. Escalating forest fire risk threatens catastrophic carbon losses in the Amazon and major population health impacts from wildfire smoke in south Asia and east Africa.

Tapping into the physiological responses to mistletoe infection during heat and drought stress.

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.