Carbon Cycling In Tropical Forests Research Articles

Tropical tree species differ in damage and mortality from lightning.

Lightning is an important agent of mortality for large tropical trees with implications for tree demography and forest carbon budgets. We evaluated interspecific differences in susceptibility to lightning damage using a unique dataset of systematically located lightning strikes in central Panama. We measured differences in mortality among trees damaged by lightning and related those to damage frequency and tree functional traits. Eighteen of 30 focal species had lightning mortality rates that deviated from null expectations. Several species showed little damage and three species had no mortality from lightning, whereas palms were especially likely to die from strikes. Species that were most likely to be struck also showed the highest survival. Interspecific differences in tree tolerance to lightning suggest that lightning-caused mortality shapes compositional dynamics over time and space. Shifts in lightning frequency due to climatic change are likely to alter species composition and carbon cycling in tropical forests.

Stem Carbon Dioxide Efflux of Lignophytes Exceeds That of Cycads and Arborescent Monocots

Tree stem CO2 efflux (Es) can be substantial and the factors controlling ecosystem-level Es are required to fully understand the carbon cycle and construct models that predict atmospheric CO2 dynamics. The majority of Es studies used woody lignophyte trees as the model species. Applying these lignophyte data to represent all tree forms can be inaccurate. The Es of 318 arborescent species was quantified in a common garden setting and the results were sorted into four stem growth forms: cycads, palms, monocot trees that were not palms, and woody lignophyte trees. The woody trees were comprised of gymnosperm and eudicot species. The Es did not differ among the cycads, palms, and non-palm monocots. Lignophyte trees exhibited Es that was 40% greater than that of the other stem growth forms. The Es of lignophyte gymnosperm trees was similar to that of lignophyte eudicot trees. This extensive species survey indicates that the Es from lignophyte tree species do not align with the Es from other tree growth forms. Use of Es estimates from the literature can be inaccurate for understanding the carbon cycle in tropical forests, which contain numerous non-lignophyte tree species.

Leaf-litter production in human-modified Amazonian forests following the El Niño-mediated drought and fires of 2015–2016

Leaf-litter production is an essential part of the carbon cycle of tropical forests. In the Amazon, it is influenced by climate, presenting high levels during the driest months of the year. However, it is less established how extreme climatic events may impact leaf-litter production in the long term. Even more unclear is how litter production is affected by human-driven disturbances. Here we examine the effects of the 2015–16 El Niño drought and subsequent fires in the leaf-litter production of human-modified Amazonian forests, thus investigating the interactions of a climatic extreme with anthropogenic disturbances on this key process of the Amazonian carbon cycle. We sampled leaf litter from April 2015 until March 2019 across 20 plots located in the eastern Brazilian Amazon, in a total of 11,548 samples. Plots were distributed along a pre-El Niño gradient of human disturbance, including undisturbed, logged, logged-and-burned, and secondary forests. All plots were impacted by the extreme drought caused by the 2015–16 El Niño, and eight were also impacted by understory fires. We found a significant and non-linear relationship between precipitation and monthly leaf-litter production – above 300 mm of monthly precipitation, the production of leaf-litter becomes independent of rainfall. Surprisingly, this relationship was not influenced by pre-El Niño forest disturbance class. During the El Niño, leaf-litter production was higher, decreasing sharply in the following year, especially in El Niño-fire-affected forests. Between 2017 and 2019, all forests experienced a gradual increase in the production of leaf litter. However, the mechanisms behind this increase remain unclear and are likely different between forests affected only by the El Niño drought and those affected by both the drought and fires. Our results suggest that while leaf-litter production may be insensitive to past human disturbances, it is affected, in the short term, by extreme climatic events, especially in forests impacted by El Niño fires.

Implications of size-dependent tree mortality for tropical forest carbon dynamics.

Tropical forests are mitigating the ongoing climate crisis by absorbing more atmospheric carbon than they emit. However, widespread increases in tree mortality rates are decreasing the ability of tropical forests to assimilate and store carbon. A relatively small number of large trees dominate the contributions of these forests to the global carbon budget, yet we know remarkably little about how these large trees die. Here, we propose a cohesive and empirically informed framework for understanding and investigating size-dependent drivers of tree mortality. This theory-based framework enables us to posit that abiotic drivers of tree mortality-particularly drought, wind and lightning-regulate tropical forest carbon cycling via their disproportionate effects on large trees. As global change is predicted to increase the pressure from abiotic drivers, the associated deaths of large trees could rapidly and lastingly reduce tropical forest biomass stocks. Focused investigations of large tree death are needed to understand how shifting drivers of mortality are restructuring carbon cycling in tropical forests.

Climate implications on forest above- and belowground carbon allocation patterns along a tropical elevation gradient on Mt. Kilimanjaro (Tanzania)

Tropical forests represent the largest store of terrestrial biomass carbon (C) on earth and contribute over-proportionally to global terrestrial net primary productivity (NPP). How climate change is affecting NPP and C allocation to tree components in forests is not well understood. This is true for tropical forests, but particularly for African tropical forests. Studying forest ecosystems along elevation and related temperature and moisture gradients is one possible approach to address this question. However, the inclusion of belowground productivity data in such studies is scarce. On Mt. Kilimanjaro (Tanzania), we studied aboveground (wood increment, litter fall) and belowground (fine and coarse root) NPP along three elevation transects (c. 1800–3900 m a.s.l.) across four tropical montane forest types to derive C allocation to the major tree components. Total NPP declined continuously with elevation from 8.5 to 2.8 Mg C ha−1 year−1 due to significant decline in aboveground NPP, while fine root productivity (sequential coring approach) remained unvaried with around 2 Mg C ha−1 year−1, indicating a marked shift in C allocation to belowground components with elevation. The C and N fluxes to the soil via root litter were far more important than leaf litter inputs in the subalpine Erica forest. Thus, the shift of C allocation to belowground organs with elevation at Mt. Kilimanjaro and other tropical forests suggests increasing nitrogen limitation of aboveground tree growth at higher elevations. Our results show that studying fine root productivity is crucial to understand climate effects on the carbon cycle in tropical forests.

Fungal succession in decomposing woody debris across a tropical forest disturbance gradient

Fungi decompose woody debris, an important carbon pool in forests. Fungal community structure is expected to vary according to the wood species, habitats and extent of abiotic disturbance, which have consequences for carbon cycling in tropical forests. Here we examined the effects of fungal diversity and composition on woody debris decomposition rates and sought potential mechanisms to explain an observed lack of difference in decomposition rates across a disturbance gradient in a tropical montane rainforest in Xishuangbanna, SW China. We measured wood specific gravity (WSG) loss from 280 logs of Litsea cubeba and Castanopsis mekongensis over 3 years and monitored fungal communities from 418 samples using next-generation sequencing after 0, 18 and 36 months field exposure. Wood species and termite presence determined changes in fungal diversity through time. Overall there was a peak in fungal diversity at 18 mo, suggesting an initial period of colonization followed by a period of increasingly competitive interactions leading to decreased diversity. Litsea logs, which had relatively low initial WSG and thinner bark, harbored higher fungal diversity. Shared fungal OTUs between wood species peaked at 18 mo (~50%). However, fungal diversity was not a significant predictor of WSG loss. An effect of habitat on fungal community composition suggests that functional replacement explains the similar decay rates across the disturbance gradient. In addition, the proportions of saprotroph and white-rot fungi increased through time regardless of wood species. Termite presence reduced WSG loss, but the effect was mediated via the abundance of soft rot fungi. Our results suggest that changes in functional traits, rather than fungal species diversity, may better explain variation in WSG loss. Future studies should investigate roles of fungal functional traits and rot types, particularly those of Ascomycete fungi, whose roles in wood decay are still poorly characterized.

A mechanistic and empirically supported lightning risk model for forest trees

Abstract Tree death due to lightning influences tropical forest carbon cycling and tree community dynamics. However, the distribution of lightning damage among trees in forests remains poorly understood. We developed models to predict direct and secondary lightning damage to trees based on tree size, crown exposure and local forest structure. We parameterized these models using data on the locations of lightning strikes and censuses of tree damage in strike zones, combined with drone‐based maps of tree crowns and censuses of all trees within a 50‐ha forest dynamics plot on Barro Colorado Island, Panama. The likelihood of a direct strike to a tree increased with larger exposed crown area and higher relative canopy position (emergent > canopy >>> subcanopy), whereas the likelihood of secondary lightning damage increased with tree diameter and proximity to neighbouring trees. The predicted frequency of lightning damage in this mature forest was greater for tree species with larger average diameters. These patterns suggest that lightning influences forest structure and the global carbon budget by non‐randomly damaging large trees. Moreover, these models provide a framework for investigating the ecological and evolutionary consequences of lightning disturbance in tropical forests. Synthesis. Our findings indicate that the distribution of lightning damage is stochastic at large spatial grain and relatively deterministic at smaller spatial grain (<15 m). Lightning is more likely to directly strike taller trees with large crowns and secondarily damage large neighbouring trees that are closest to the directly struck tree. The results provide a framework for understanding how lightning can affect forest structure, forest dynamics and carbon cycling. The resulting lightning risk model will facilitate informed investigations into the effects of lightning in tropical forests.

Focus on the role of forests and soils in meeting climate change mitigation goals: summary

It is clear that reducing greenhouse gas emissions alone is insufficient to avoid large global temperature increases. To avoid atmospheric concentrations of greenhouse gases that result in dangerous alterations of the climate, large reductions in carbon dioxide emissions from fossil fuel combustion and land use changes must be accompanied by an increase in atmospheric carbon dioxide sequestration. Natural Climate Solutions have become a major focus of climate policy. Land and ocean ecosystems remove and store atmospheric carbon, and forests play a major role. This focus collection includes papers that address three important aspects of the role for forests in meeting climate change mitigation goals: (i) Carbon Accounting of forest sinks and reservoirs, process emissions and carbon storage in forest products, (ii) the carbon dioxide dynamics of using Forest Bioenergy and (iii) the carbon cycle of Tropical Forests.

Modeling the impact of liana infestation on the demography and carbon cycle of tropical forests.

There is mounting empirical evidence that lianas affect the carbon cycle of tropical forests. However, no single vegetation model takes into account this growth form, although such efforts could greatly improve the predictions of carbon dynamics in tropical forests. In this study, we incorporated a novel mechanistic representation of lianas in a dynamic global vegetation model (the Ecosystem Demography Model). We developed a liana‐specific plant functional type and mechanisms representing liana–tree interactions (such as light competition, liana‐specific allometries, and attachment to host trees) and parameterized them according to a comprehensive literature meta‐analysis. We tested the model for an old‐growth forest (Paracou, French Guiana) and a secondary forest (Gigante Peninsula, Panama). The resulting model simulations captured many features of the two forests characterized by different levels of liana infestation as revealed by a systematic comparison of the model outputs with empirical data, including local census data from forest inventories, eddy flux tower data, and terrestrial laser scanner‐derived forest vertical structure. The inclusion of lianas in the simulations reduced the secondary forest net productivity by up to 0.46 tC ha−1 year−1, which corresponds to a limited relative reduction of 2.6% in comparison with a reference simulation without lianas. However, this resulted in significantly reduced accumulated above‐ground biomass after 70 years of regrowth by up to 20 tC/ha (19% of the reference simulation). Ultimately, the simulated negative impact of lianas on the total biomass was almost completely cancelled out when the forest reached an old‐growth successional stage. Our findings suggest that lianas negatively influence the forest potential carbon sink strength, especially for young, disturbed, liana‐rich sites. In light of the critical role that lianas play in the profound changes currently experienced by tropical forests, this new model provides a robust numerical tool to forecast the impact of lianas on tropical forest carbon sinks.

Changes in seasonal precipitation distribution but not annual amount affect litter decomposition in a secondary tropical forest.

In the tropics of South China, climate change induced more rainfall events in the wet season in the last decades. Moreover, there will be more frequently spring drought in the future. However, knowledge on how litter decomposition rate would respond to these seasonal precipitation changes is still limited. In the present study, we conducted a precipitation manipulation experiment in a tropical forest. First, we applied a 60% rainfall exclusion in April and May to defer the onset of wet season and added the same amount of water in October and November to mimic a deferred wet season (DW); second, we increased as much as 25% mean annual precipitation into plots in July and August to simulate a wetter wet season (WW). Five single‐species litters, with their carbon to nitrogen ratio ranged from 27 to 49, and a mixed litter were used to explore how the precipitation change treatments would affect litter decomposition rate. The interaction between precipitation changes and litter species was not significant. The DW treatment marginally accelerated litter decomposition across six litter types. Detailed analysis showed that DW increased litter decomposition rate in the periods of January to March and October to December, when soil moisture was increased by the water addition in the dry season. In contrast, WW did not significantly affect litter decomposition rate, which was consistent with the unchanged soil moisture pattern. In conclusion, the study indicated that regardless of litter types or litter quality, the projected deferred wet season would increase litter decomposition rate, whereas the wetter wet season would not affect litter decomposition rate in the tropical forests. This study improves our knowledge of how tropical forest carbon cycling in response to precipitation change.

Leaf reflectance spectroscopy captures variation in carboxylation capacity across species, canopy environment and leaf age in lowland moist tropical forests.

Understanding the pronounced seasonal and spatial variation in leaf carboxylation capacity (Vc,max ) is critical for determining terrestrial carbon cycling in tropical forests. However, an efficient and scalable approach for predicting Vc,max is still lacking. Here the ability of leaf spectroscopy for rapid estimation of Vc,max was tested. Vc,max was estimated using traditional gas exchange methods, and measured reflectance spectra and leaf age in leaves sampled from tropical forests in Panama and Brazil. These data were used to build a model to predict Vc,max from leaf spectra. The results demonstrated that leaf spectroscopy accurately predicts Vc,max of mature leaves in Panamanian tropical forests (R2 =0.90). However, this single-age model required recalibration when applied to broader leaf demographic classes (i.e. immature leaves). Combined use of spectroscopy models for Vc,max and leaf age enabled construction of the Vc,max -age relationship solely from leaf spectra, which agreed with field observations. This suggests that the spectroscopy technique can capture the seasonal variability in Vc,max , assuming sufficient sampling across diverse species, leaf ages and canopy environments. This finding will aid development of remote sensing approaches that can be used to characterize Vc,max in moist tropical forests and enable an efficient means to parameterize and evaluate terrestrial biosphere models.

Forest biomass, productivity and carbon cycling along a rainfall gradient in West Africa.

Net Primary Productivity (NPP) is one of the most important parameters in describing the functioning of any ecosystem and yet it arguably remains a poorly quantified and understood component of carbon cycling in tropical forests, especially outside of the Americas. We provide the first comprehensive analysis of NPP and its carbon allocation to woody, canopy and root growth components at contrasting lowland West African forests spanning a rainfall gradient. Using a standardized methodology to study evergreen (EF), semi-deciduous (SDF), dry forests (DF) and woody savanna (WS), we find that (i) climate is more closely related with above and belowground C stocks than with NPP (ii) total NPP is highest in the SDF site, then the EF followed by the DF and WS and that (iii) different forest types have distinct carbon allocation patterns whereby SDF allocate in excess of 50% to canopy production and the DF and WS sites allocate 40%-50% to woody production. Furthermore, we find that (iv) compared with canopy and root growth rates the woody growth rate of these forests is a poor proxy for their overall productivity and that (v) residence time is the primary driver in the productivity-allocation-turnover chain for the observed spatial differences in woody, leaf and root biomass across the rainfall gradient. Through a systematic assessment of forest productivity we demonstrate the importance of directly measuring the main components of above and belowground NPP and encourage the establishment of more permanent carbon intensive monitoring plots across the tropics.

Nutrient limitations to bacterial and fungal growth during cellulose decomposition in tropical forest soils

Nutrients constrain the soil carbon cycle in tropical forests, but we lack knowledge on how these constraints vary within the soil microbial community. Here, we used in situ fertilization in a montane tropical forest and in two lowland tropical forests on contrasting soil types to test the principal hypothesis that there are different nutrient constraints to different groups of microorganisms during the decomposition of cellulose. We also tested the hypotheses that decomposers shift from nitrogen to phosphorus constraints from montane to lowland forests, respectively, and are further constrained by potassium and sodium deficiency in the western Amazon. Cellulose and nutrients (nitrogen, phosphorus, potassium, sodium, and combined) were added to soils in situ, and microbial growth on cellulose (phospholipid fatty acids and ergosterol) and respiration were measured. Microbial growth on cellulose after single nutrient additions was highest following nitrogen addition for fungi, suggesting nitrogen as the primary limiting nutrient for cellulose decomposition. This was observed at all sites, with no clear shift in nutrient constraints to decomposition between lowland and montane sites. We also observed positive respiration and fungal growth responses to sodium and potassium addition at one of the lowland sites. However, when phosphorus was added, and especially when added in combination with other nutrients, bacterial growth was highest, suggesting that bacteria out-compete fungi for nitrogen where phosphorus is abundant. In summary, nitrogen constrains fungal growth and cellulose decomposition in both lowland and montane tropical forest soils, but additional nutrients may also be of critical importance in determining the balance between fungal and bacterial decomposition of cellulose.

Role of tree size in moist tropical forest carbon cycling and water deficit responses.

Drought disproportionately affects larger trees in tropical forests, but implications for forest composition and carbon (C) cycling in relation to dry season intensity remain poorly understood. In order to characterize how C cycling is shaped by tree size and drought adaptations and how these patterns relate to spatial and temporal variation in water deficit, we analyze data from three forest dynamics plots spanning a moisture gradient in Panama that have experienced El Niño droughts. At all sites, aboveground C cycle contributions peaked below 50-cm stem diameter, with stems ≥50cm accounting for on average 59% of live aboveground biomass, 45% of woody productivity and 49% of woody mortality. The dominance of drought-avoidance strategies increased interactively with stem diameter and dry season intensity. Although size-related C cycle contributions did not vary systematically across the moisture gradient under nondrought conditions, woody mortality of larger trees was disproportionately elevated under El Niño drought stress. Thus, large (>50cm) stems, which strongly mediate but do not necessarily dominate C cycling, have drought adaptations that compensate for their more challenging hydraulic environment, particularly in drier climates. However, these adaptations do not fully buffer the effects of severe drought, and increased large tree mortality dominates ecosystem-level drought responses.

Temperature and rainfall interact to control carbon cycling in tropical forests.

Tropical forests dominate global terrestrial carbon (C) exchange, and recent droughts in the Amazon Basin have contributed to short-term declines in terrestrial carbon dioxide uptake and storage. However, the effects of longer-term climate variability on tropical forest carbon dynamics are still not well understood. We synthesised field data from more than 150 tropical forest sites to explore how climate regulates tropical forest aboveground net primary productivity (ANPP) and organic matter decomposition, and combined those data with two existing databases to explore climate - C relationships globally. While previous analyses have focused on the effects of either temperature or rainfall on ANPP, our results highlight the importance of interactions between temperature and rainfall on the C cycle. In cool forests (<20°C), high rainfall slowed rates of C cycling, but in warm tropical forests (>20°C) it consistently enhanced both ANPP and decomposition. At the global scale, our analysis showed an increase in ANPP with rainfall in relatively warm sites, inconsistent with declines in ANPP with rainfall reported previously. Overall, our results alter our understanding of climate - C cycle relationships, with high precipitation accelerating rates of C exchange with the atmosphere in the most productive biome on earth.

Trailblazing the Carbon Cycle of Tropical Forests from Puerto Rico

We review the literature that led to clarifying the role of tropical forests in the global carbon cycle from a time when they were considered sources of atmospheric carbon to the time when they were found to be atmospheric carbon sinks. This literature originates from work conducted by US Forest Service scientists in Puerto Rico and their collaborators. It involves the classification of forests by life zones, estimation of carbon density by forest type, assessing carbon storage changes with ecological succession and land use/land cover type, describing the details of the carbon cycle of forests at stand and landscape levels, assessing global land cover by forest type and the complexity of land use change in tropical regions, and assessing the ecological fluxes and storages that contribute to net carbon accumulation in tropical forests. We also review recent work that couples field inventory data, remote sensing technology such as LIDAR, and GIS analysis in order to more accurately determine the role of tropical forests in the global carbon cycle and point out new avenues of carbon research that address the responses of tropical forests to environmental change.

Climate seasonality limits leaf carbon assimilation and wood productivity in tropical forests

Abstract. The seasonal climate drivers of the carbon cycle in tropical forests remain poorly known, although these forests account for more carbon assimilation and storage than any other terrestrial ecosystem. Based on a unique combination of seasonal pan-tropical data sets from 89 experimental sites (68 include aboveground wood productivity measurements and 35 litter productivity measurements), their associated canopy photosynthetic capacity (enhanced vegetation index, EVI) and climate, we ask how carbon assimilation and aboveground allocation are related to climate seasonality in tropical forests and how they interact in the seasonal carbon cycle. We found that canopy photosynthetic capacity seasonality responds positively to precipitation when rainfall is &lt; 2000 mm yr−1 (water-limited forests) and to radiation otherwise (light-limited forests). On the other hand, independent of climate limitations, wood productivity and litterfall are driven by seasonal variation in precipitation and evapotranspiration, respectively. Consequently, light-limited forests present an asynchronism between canopy photosynthetic capacity and wood productivity. First-order control by precipitation likely indicates a decrease in tropical forest productivity in a drier climate in water-limited forest, and in current light-limited forest with future rainfall &lt; 2000 mm yr−1.

Source and sink carbon dynamics and carbon allocation in the Amazon basin

AbstractChanges to the carbon cycle in tropical forests could affect global climate, but predicting such changes has been previously limited by lack of field‐based data. Here we show seasonal cycles of the complete carbon cycle for 14, 1 ha intensive carbon cycling plots which we separate into three regions: humid lowland, highlands, and dry lowlands. Our data highlight three trends: (1) there is differing seasonality of total net primary productivity (NPP) with the highlands and dry lowlands peaking in the dry season and the humid lowland sites peaking in the wet season, (2) seasonal reductions in wood NPP are not driven by reductions in total NPP but by carbon during the dry season being preferentially allocated toward either roots or canopy NPP, and (3) there is a temporal decoupling between total photosynthesis and total carbon usage (plant carbon expenditure). This decoupling indicates the presence of nonstructural carbohydrates which may allow growth and carbon to be allocated when it is most ecologically beneficial rather than when it is most environmentally available.

The linkages between photosynthesis, productivity, growth and biomass in lowland Amazonian forests

Understanding the relationship between photosynthesis, net primary productivity and growth in forest ecosystems is key to understanding how these ecosystems will respond to global anthropogenic change, yet the linkages among these components are rarely explored in detail. We provide the first comprehensive description of the productivity, respiration and carbon allocation of contrasting lowland Amazonian forests spanning gradients in seasonal water deficit and soil fertility. Using the largest data set assembled to date, ten sites in three countries all studied with a standardized methodology, we find that (i) gross primary productivity (GPP) has a simple relationship with seasonal water deficit, but that (ii) site-to-site variations in GPP have little power in explaining site-to-site spatial variations in net primary productivity (NPP) or growth because of concomitant changes in carbon use efficiency (CUE), and conversely, the woody growth rate of a tropical forest is a very poor proxy for its productivity. Moreover, (iii) spatial patterns of biomass are much more driven by patterns of residence times (i.e. tree mortality rates) than by spatial variation in productivity or tree growth. Current theory and models of tropical forest carbon cycling under projected scenarios of global atmospheric change can benefit from advancing beyond a focus on GPP. By improving our understanding of poorly understood processes such as CUE, NPP allocation and biomass turnover times, we can provide more complete and mechanistic approaches to linking climate and tropical forest carbon cycling.

Drought impact on forest carbon dynamics and fluxes in Amazonia.

In 2005 and 2010 the Amazon basin experienced two strong droughts, driven by shifts in the tropical hydrological regime possibly associated with global climate change, as predicted by some global models. Tree mortality increased after the 2005 drought, and regional atmospheric inversion modelling showed basin-wide decreases in CO2 uptake in 2010 compared with 2011 (ref. 5). But the response of tropical forest carbon cycling to these droughts is not fully understood and there has been no detailed multi-site investigation in situ. Here we use several years of data from a network of thirteen 1-ha forest plots spread throughout South America, where each component of net primary production (NPP), autotrophic respiration and heterotrophic respiration is measured separately, to develop a better mechanistic understanding of the impact of the 2010 drought on the Amazon forest. We find that total NPP remained constant throughout the drought. However, towards the end of the drought, autotrophic respiration, especially in roots and stems, declined significantly compared with measurements in 2009 made in the absence of drought, with extended decreases in autotrophic respiration in the three driest plots. In the year after the drought, total NPP remained constant but the allocation of carbon shifted towards canopy NPP and away from fine-root NPP. Both leaf-level and plot-level measurements indicate that severe drought suppresses photosynthesis. Scaling these measurements to the entire Amazon basin with rainfall data, we estimate that drought suppressed Amazon-wide photosynthesis in 2010 by 0.38petagrams of carbon (0.23-0.53petagrams of carbon). Overall, we find that during this drought, instead of reducing total NPP, trees prioritized growth by reducing autotrophic respiration that was unrelated to growth. This suggests that trees decrease investment in tissue maintenance and defence, in line with eco-evolutionary theories that trees are competitively disadvantaged in the absence of growth. We propose that weakened maintenance and defence investment may, in turn, cause the increase in post-drought tree mortality observed at our plots.