Thirty-eight years of CO<sub>2</sub> fertilization has outpaced growing aridity to drive greening of Australian woody ecosystems

Climate change is projected to increase the imbalance between the supply (precipitation) and atmospheric demand for water (i.e., increased potential evapotranspiration), stressing plants in water-limited environments. Plants may be able to offset increasing aridity because rising CO2 increases water use efficiency. CO2 fertilization has also been cited as one of the drivers of the widespread "greening" phenomenon. However, attributing the size of this CO2 fertilization effect is complicated, due in part to a lack of long-term vegetation monitoring and interannual- to decadalscale climate variability. In this study we asked the question of how much CO2 has contributed towards greening. We focused our analysis on a broad aridity gradient spanning eastern Australia's woody ecosystems. Next we analyzed 38 years of satellite remote sensing estimates of vegetation greenness (normalized difference vegetation index, NDVI) to examine the role of CO2 in ameliorating climate change impacts. Multiple statistical techniques were applied to separate the CO2-attributable effects on greening from the changes in water supply and atmospheric aridity. Widespread vegetation greening occurred despite a warming climate, increases in vapor pressure deficit, and repeated record-breaking droughts and heat waves. Between 1982-2019 we found that NDVI increased (median 11.3 %) across 90.5 % of the woody regions. After masking disturbance effects (e.g., fire), we statistically estimated an 11.7 % increase in NDVI attributable to CO2, broadly consistent with a hypothesized theoretical expectation of an 8.6 % increase in water use efficiency due to rising CO2. In contrast to reports of a weakening CO2 fertilization effect, we found no consistent temporal change in the CO2 effect. We conclude rising CO2 has mitigated the effects of increasing aridity, repeated record-breaking droughts, and record-breaking heat waves in eastern Australia. However, we were unable to determine whether trees or grasses were the primary beneficiary of the CO2-induced change in water use efficiency, which has implications for projecting future ecosystem resilience. A more complete understanding of how CO2-induced changes in water use efficiency affect trees and non-tree vegetation is needed

Phenology and Seasonal Ecosystem Productivity in an Amazonian Floodplain Forest

everal studies have explored the linkages between phenology and ecosystem productivity across the Amazon basin. However, few studies have focused on flooded forests, which correspond to c.a. 14% of the basin. In this study, we assessed the seasonality of ecosystem productivity (gross primary productivity, GPP) from eddy covariance measurements, environmental drivers and phenological patterns obtained from the field (leaf litter mass) and satellite measurements (enhanced vegetation index (EVI) from the Moderate Resolution Imaging Spectroradiometer/multi-angle implementation correction (MODIS/MAIAC)) in an Amazonian floodplain forest. We found that ecosystem productivity is limited by soil moisture in two different ways. During the flooded period, the excess of water limits GPP (Spearman’s correlation; rho = −0.22), while during non-flooded months, GPP is positively associated with soil moisture (rho = 0.34). However, GPP is maximized when cumulative water deficit (CWD) increases (rho = 0.81), indicating that GPP is dependent on the amount of water available. EVI was positively associated with leaf litter mass (Pearson’s correlation; r = 0.55) and with GPP (r = 0.50), suggesting a coupling between new leaf production and the phenology of photosynthetic capacity, decreasing both at the peak of the flooded period and at the end of the dry season. EVI was able to describe the inter-annual variations on forest responses to environmental drivers, which have changed during an observed El Niño-Southern Oscillation (ENSO) year (2015/2016)

Towards species-level forecasts of drought-induced tree mortality risk

Predicting species-level responses to drought at the landscape scale is critical to reducing uncertainty in future terrestrial carbon and water cycle projections. We embedded a stomatal optimisation model in the Community Atmosphere Biosphere Land Exchange (CABLE) land surface model and parameterised the model for 15 canopy dominant eucalypt tree species across South-Eastern Australia (mean annual precipitation range: 344–1424 mm yr−1). We conducted three experiments: applying CABLE to the 2017–2019 drought; a 20% drier drought; and a 20% drier drought with a doubling of atmospheric carbon dioxide (CO2). The severity of the drought was highlighted as for at least 25% of their distribution ranges, 60% of species experienced leaf water potentials beyond the water potential at which 50% of hydraulic conductivity is lost due to embolism. We identified areas of severe hydraulic stress within-species’ ranges, but we also pinpointed resilience in species found in predominantly semiarid areas. The importance of the role of CO2 in ameliorating drought stress was consistent across species. Our results represent an important advance in our capacity to forecast the resilience of individual tree species, providing an evidence base for decision-making around the resilience of restoration plantings or net-zero emission strategies

Towards species‐level forecasts of drought‐induced tree mortality risk

Predicting species-level responses to drought at the landscape scale is critical to reducing uncertainty in future terrestrial carbon and water cycle projections. We embedded a stomatal optimisation model in the Community Atmosphere Biosphere Land Exchange (CABLE) land surface model and parameterised the model for 15 canopy dominant eucalypt tree species across South-Eastern Australia (mean annual precipitation range: 344–1424 mm yr−1). We conducted three experiments: applying CABLE to the 2017–2019 drought; a 20% drier drought; and a 20% drier drought with a doubling of atmospheric carbon dioxide (CO2). The severity of the drought was highlighted as for at least 25% of their distribution ranges, 60% of species experienced leaf water potentials beyond the water potential at which 50% of hydraulic conductivity is lost due to embolism. We identified areas of severe hydraulic stress within-species’ ranges, but we also pinpointed resilience in species found in predominantly semiarid areas. The importance of the role of CO2 in ameliorating drought stress was consistent across species. Our results represent an important advance in our capacity to forecast the resilience of individual tree species, providing an evidence base for decision-making around the resilience of restoration plantings or net-zero emission strategies

The structures underpinning vulnerability: examining landscape-society interactions in a smallholder coffee agroforestry system

Smallholder farmers dependent on rain-fed agriculture are particularly vulnerable to extreme climate events and, therefore, it is necessary to identify adaptive measures that would increase farmer resilience to these shocks. The management options in a low-input system, like forest coffee (Coffea arabica), are limited and there are several factors out of farmers' control driving their vulnerability to changing climatic conditions. These can relate to social structures and landscape factors, which can interact to reduce farmers' adaptive capacity, creating a state of contextual vulnerability. We explored the potential synergies of this interaction across elevation, patch area and shade management gradients for smallholder coffee farms around the UNESCO Yayu Coffee Forest Biosphere Reserve in Ethiopia before, during and immediately following the 2015/16 El Niño. We documented a dramatic collapse in coffee yields across all farms, resulting in coffee incomes 29.5 ± 18.0 % and 19.5 ± 10.0 % of 2014 incomes in 2015 and 2016, respectively. We identified farms at elevations between 1500-1600 m with canopy openness between 40-45 % as being consistently low yielding over our study period. We found these farmers had the highest rates of income diversification and, therefore, were already exhibiting adaptive capacity. Farmers with the largest income losses were spatially concentrated between 1600-1700 m, located in larger patch areas with lower canopy openness. Farmers at this elevation have access to poor infrastructure, restrictions on shade management and reported higher dependence on income from coffee, indicating an interaction of biotic and social factors exacerbating their vulnerability. Unfortunately, due to a nationally declared state of emergency, we were unable to survey farmers on the adaptive measures they undertook; therefore, we are limited in assessing their resilience. However, we do show the importance of considering both biotically and socially-mediated influences for assessing smallholder vulnerability, particularly barriers to diversifying incomes

Identifying areas at risk of drought-induced tree mortality across South-Eastern Australia

South-East Australia has recently been subjected to two of the worst droughts in the historical record (Millennium Drought, 2000–2009 and Big Dry, 2017–2019). Unfortunately, a lack of forest monitoring has made it difficult to determine whether widespread tree mortality has resulted from these droughts. Anecdotal observations suggest the Big Dry may have led to more significant tree mortality than the Millennium drought. Critically, to be able to robustly project future expected climate change effects on Australian vegetation, we need to be able to assess the vulnerability to drought of Australian trees. Here, we implemented a model of plant hydraulics into the Community Atmosphere Biosphere Land Exchange (CABLE) land surface model. We parameterised the drought response behaviour of five broad vegetation types, based on a common garden dry-down experiment with species originating across a rainfall gradient (188–1125 mm yr1 ) across South-East Australia. The new hydraulics model significantly improved (~35–45 % reduction in root mean square error) CABLE’s previous predictions of latent heat fluxes during periods of water stress at two eddy covariance sites in Australia. Landscape-scale predictions of the greatest percentage loss of hydraulic conductivity (PLC), 40–60 %, were broadly consistent with satellite estimates of regions of the greatest change in both droughts. In neither drought did CABLE predict that trees would have reached critical PLC in widespread areas (i.e. it projected a low mortality risk), although the model highlighted critical levels near the desert regions of South-East Australia where few trees live. Overall, our experimentally constrained model results imply significant resilience to drought conferred by hydraulic function, but also highlight critical data and scientific gaps. Our approach presents a promising avenue to integrate experimental data and make regional-scale predictions of potential drought-induced hydraulic failure

Fine root dynamics across pantropical rainforest ecosystems

Fine roots constitute a significant component of the net primary productivity (NPP) of forest ecosystems but are much less studied than aboveground NPP. Comparisons across sites and regions are also hampered by inconsistent methodologies, especially in tropical areas. Here, we present a novel dataset of fine root biomass, productivity, residence time, and allocation in tropical old-growth rainforest sites worldwide, measured using consistent methods, and examine how these variables are related to consistently determined soil and climatic characteristics. Our pantropical dataset spans intensive monitoring plots in lowland (wet, semi-deciduous, and deciduous) and montane tropical forests in South America, Africa, and Southeast Asia (n = 47). Large spatial variation in fine root dynamics was observed across montane and lowland forest types. In lowland forests, we found a strong positive linear relationship between fine root productivity and sand content, this relationship was even stronger when we considered the fractional allocation of total NPP to fine roots, demonstrating that understanding allocation adds explanatory power to understanding fine root productivity and total NPP. Fine root residence time was a function of multiple factors: soil sand content, soil pH, and maximum water deficit, with longest residence times in acidic, sandy, and water-stressed soils. In tropical montane forests, on the other hand, a different set of relationships prevailed, highlighting the very different nature of montane and lowland forest biomes. Root productivity was a strong positive linear function of mean annual temperature, root residence time was a strong positive function of soil nitrogen content in montane forests, and lastly decreasing soil P content increased allocation of productivity to fine roots. In contrast to the lowlands, environmental conditions were a better predictor for fine root productivity than for fractional allocation of total NPP to fine roots, suggesting that root productivity is a particularly strong driver of NPP allocation in tropical mountain regions.WHH was funded by Peruvian FONDECYT/CONCYTEC (grant contract number 213-2015-FONDECYT). The GEM network was supported by a European Research Council Advanced Investigator Grant to YM (GEM-TRAITS: 321131) under the European Union's Seventh Framework Programme (FP7/2007-2013). The field data collection was funded NERC Grants NE/D014174/1 and NE/J022616/1 for in Peru, BALI (NE/K016369/1) for work in Malaysia, the Royal Society-Leverhulme Africa Capacity Building Programme for work in Ghana and Gabon and ESPA-ECOLIMITS (NE/1014705/1) in Ghana and Ethiopia. Plot inventories in South America were supported by funding from the US National Science Foundation Long-Term Research in Environmental Biology program (LTREB; DEB 1754647) and the Gordon and Betty Moore Foundation Andes-Amazon Program. GEM data in Gabon were collected under authorization to YM and supported by the Gabon National Parks Agency. Y.M. is supported by the Jackson Foundation. We would like to acknowledge the GEM team across the tropical regions and countries of Bolivia, Brazil, Ghana, Gabon, Ethiopia, Malaysia, and Peru

Windthrows control biomass patterns and functional composition of Amazon forests

Amazon forests account for ~25% of global land biomass and tropical tree species. In these forests, windthrows (i.e., snapped and uprooted trees) are a major natural disturbance, but the rates and mechanisms of recovery are not known. To provide a predictive framework for understanding the effects of windthrows on forest structure and functional composition (DBH ≥10 cm), we quantified biomass recovery as a function of windthrow severity (i.e., fraction of windthrow tree mortality on Landsat pixels, ranging from 0%–70%) and time since disturbance for terra-firme forests in the Central Amazon. Forest monitoring allowed insights into the processes and mechanisms driving the net biomass change (i.e., increment minus loss) and shifts in functional composition. Windthrown areas recovering for between 4–27 years had biomass stocks as low as 65.2–91.7 Mg/ha or 23%–38% of those in nearby undisturbed forests (~255.6 Mg/ha, all sites). Even low windthrow severities (4%–20% tree mortality) caused decadal changes in biomass stocks and structure. While rates of biomass increment in recovering vegetation were nearly double (6.3 ± 1.4 Mg ha− 1 year− 1) those of undisturbed forests (~3.7 Mg ha− 1 year− 1), biomass loss due to post-windthrow mortality was high (up to −7.5 ± 8.7 Mg ha− 1 year− 1, 8.5 years since disturbance) and unpredictable. Consequently, recovery to 90% of “pre-disturbance” biomass takes up to 40 years. Resprouting trees contributed little to biomass recovery. Instead, light-demanding, low-density genera (e.g., Cecropia, Inga, Miconia, Pourouma, Tachigali, and Tapirira) were favored, resulting in substantial post-windthrow species turnover. Shifts in functional composition demonstrate that windthrows affect the resilience of live tree biomass by favoring soft-wooded species with shorter life spans that are more vulnerable to future disturbances. As the time required for forests to recover biomass is likely similar to the recurrence interval of windthrows triggering succession, windthrows have the potential to control landscape biomass/carbon dynamics and functional composition in Amazon forests. ©2018 The Authors. Global Change Biology Published by John Wiley & Sons Lt

Vulnerability of Amazon forests to storm-driven tree mortality

Tree mortality is a key driver of forest community composition and carbon dynamics. Strong winds associated with severe convective storms are dominant natural drivers of tree mortality in the Amazon. Why forests vary with respect to their vulnerability to wind events and how the predicted increase in storm events might affect forest ecosystems within the Amazon are not well understood. We found that windthrows are common in the Amazon region extending from northwest (Peru, Colombia, Venezuela, and west Brazil) to central Brazil, with the highest occurrence of windthrows in the northwest Amazon. More frequent winds, produced by more frequent severe convective systems, in combination with well-known processes that limit the anchoring of trees in the soil, help to explain the higher vulnerability of the northwest Amazon forests to winds. Projected increases in the frequency and intensity of convective storms in the Amazon have the potential to increase wind-related tree mortality. A forest demographic model calibrated for the northwestern and the central Amazon showed that northwestern forests are more resilient to increased wind-related tree mortality than forests in the central Amazon. Our study emphasizes the importance of including wind-related tree mortality in model simulations for reliable predictions of the future of tropical forests and their effects on the Earth' system