See Letter to the Editor on this article by Adam et al., 26, 3756–3758. See also Response to the Letter by Bradstock et al., 26, e8–e9.

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Atmospheric carbon dioxide enrichment (eCO 2) can enhance plant carbon uptake and growth 1,2,3,4,5 , thereby providing an important negative feedback to climate change by slowing the rate of increase of the atmospheric CO 2 concentration 6. While evidence gathered from young aggrading forests has generally indicated a strong CO 2 fertilization effect on biomass growth 3,4,5 , it is unclear whether mature forests respond to eCO 2 in a similar way. In mature trees and forest stands 7,8,9,10 , photosynthetic uptake has been found to increase under eCO 2 without any apparent accompanying growth response, leaving an open question about the fate of additional carbon fixed under eCO 2 4,5,7,8,9,10,11. Here, using data from the first ecosystemscale Free-Air CO 2 Enrichment (FACE) experiment in a mature forest, we constructed a comprehensive ecosystem carbon budget to track the fate of carbon as the forest responds to four years of eCO 2 exposure. We show that, although the eCO 2 treatment of ambient +150 ppm (+38%) induced a 12% (+247 g C m-2 yr-1) 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 ~50% 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 eCO 2 , and challenge the efficacy of climate mitigation strategies that rely on ubiquitous CO 2 fertilization as a driver of increased carbon sinks in global forests. Main text Globally, forests act as a large carbon sink, absorbing a significant portion of the anthropogenic CO 2 emissions 1,12 , an ecosystem service that has tremendous social and

Extreme fire seasons characterised by very large ‘mega-fires’ have demonstrably increased area burnt across forested regions globally. However, the effect of extreme fire seasons on fire severity, a measure of fire impacts on ecosystems, remains unclear. Very large wildfires burnt an unprecedented area of temperate forest, woodland and shrubland across south-eastern Australia in 2019/2020, providing an opportunity to examine the impact of extreme fires on fire severity patterns. We developed an atlas of wildfire severity across south-eastern Australia between 1988 and 2020 to test (a) whether the 2019/2020 fire season was more severe than previous fire seasons, and (b) if the proportion of high-severity fire within the burn extent (HSp) increases with wildfire size and annual area burnt. We demonstrate that the 2019/2020 wildfires in south-eastern Australia were generally greater in extent but not proportionally more severe than previous fires, owing to constant scaling between HSp and annual fire extent across the dominant dry-forest communities. However, HSp did increase with increasing annual fire extent across wet-forests and the less-common rainforest and woodland communities. The absolute area of high-severity fire in 2019/2020 (∼1.8 M ha) was larger than previously seen, accounting for ∼44% of the area burnt by high-severity fire over the past 33 years. Our results demonstrate that extreme fire seasons are a rare but defining feature of fire regimes across forested regions, owing to the disproportionate influence of mega-fires on area burnt.

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AbstractAtmospheric carbon dioxide enrichment (eCO2) can enhance plant carbon uptake and growth1,2,3,4,5, thereby providing an important negative feedback to climate change by slowing the rate of increase of the atmospheric CO2 concentration6. While evidence gathered from young aggrading forests has generally indicated a strong CO2 fertilization effect on biomass growth3,4,5, it is unclear whether mature forests respond to eCO2 in a similar way. In mature trees and forest stands7,8,9,10, photosynthetic uptake has been found to increase under eCO2 without any apparent accompanying growth response, leaving an open question about the fate of additional carbon fixed under eCO24, 5, 7,8,9,10,11. 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 responds to four years of eCO2 exposure. We show that, although the eCO2 treatment of ambient +150 ppm (+38%) induced a 12% (+247 gCm-2yr-1) 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 contributing ∼50% 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 CO2 fertilization as a driver of increased carbon sinks in standing forests and afforestation projects.

Wildfire refugia (unburnt patches within large wildfires) are important for the persistence of fire‐sensitive species across forested landscapes globally. A key challenge is to identify the factors that determine the distribution of fire refugia across space and time. In particular, determining the relative influence of climatic and landscape factors is important in order to understand likely changes in the distribution of wildfire refugia under future climates. Here, we examine the relative effect of weather (i.e. fire weather, drought severity) and landscape features (i.e. topography, fuel age, vegetation type) on the occurrence of fire refugia across 26 large wildfires in south‐eastern Australia. Fire weather and drought severity were the primary drivers of the occurrence of fire refugia, moderating the effect of landscape attributes. Unburnt patches rarely occurred under ‘severe’ fire weather, irrespective of drought severity, topography, fuels or vegetation community. The influence of drought severity and landscape factors played out most strongly under ‘moderate’ fire weather. In mesic forests, fire refugia were linked to variables that affect fuel moisture, whereby the occurrence of unburnt patches decreased with increasing drought conditions and were associated with more mesic topographic locations (i.e. gullies, pole‐facing aspects) and vegetation communities (i.e. closed‐forest). In dry forest, the occurrence of refugia was responsive to fuel age, being associated with recently burnt areas (<5 years since fire). Overall, these results show that increased severity of fire weather and increased drought conditions, both predicted under future climate scenarios, are likely to lead to a reduction of wildfire refugia across forests of southern Australia. Protection of topographic areas able to provide long‐term fire refugia will be an important step towards maintaining the ecological integrity of forests under future climate change.

1. Wildfire occurrence and severity are projected to increase in response to anthropogenic climate change, leading to fire regimes that may exceed the limits of tolerance for some species. Plants capable of regenerating from aerial shoots following high intensity fires, termed 'epicormic resprouters', are assumed to be resilient to changes in fire regimes. However, empirical tests of the response of epicormic resprouters to extreme fire regimes, such as repeated canopy fires at short intervals, are currently lacking.2. This study examined the effect of combinations of understorey fire and canopy fire across two successive wildfires (2007, 2013) on the resilience of eucalypts that resprout epicormically. The study took place in a temperate eucalypt forest in south eastern Australia. Measures used to infer community resilience included stem topkill and damage, and seedling recruitment. It was predicted that: (a) stems will exhibit lower resistance (i.e. increased topkill and damage) to canopy fire than understorey fire; (b) recruitment will be higher following canopy fire than understorey fire; (c) prior exposure to canopy fire will reduce stem resistance and recruitment in response to subsequent wildfires; and (d) stem resistance will vary depending on bark traits.3. Topkill of saplings and small stems (<30 cm diameter at breast height) was higher in sites that recently (i.e. 2013) experienced canopy fire as opposed to understorey fire. Recent fire severity had no effect on topkill of large trees. Tree species with dense bark on the main stem and larger branches were less prone to topkill or partial stem and branch mortality than species with fibrous bark or exposed branches. Seedling recruitment was greater following canopy fire than understorey fire. Exposure to past canopy fire (i.e. in 2007) did not decrease stem resistance or recruitment. Synthesis.The results of this study suggest that communities of eucalypts that can resprout epicormically following fire will experience demographic shifts following repeated canopy fires. However, given the high resistance of large trees and rapid post-fire recovery of the seedbank, ecosystem conversion appears unlikely. The findings support the presumption that forest communities of epicormic resprouters are highly resilient to shifts in fire regimes.