Optimising height-growth predicts trait responses to water availability and other environmental drivers.

Future changes in climate, together with rising atmospheric CO 2 ${\text{CO}}_{2}$ , may reorganise the functional composition of ecosystems. Without long-term historical data, predicting how traits will respond to environmental conditions-in particular, water availability-remains a challenge. While eco-evolutionary optimality theory (EEO) can provide insight into how plants adapt to their environment, EEO approaches to date have been formulated on the assumption that plants maximise carbon gain, which omits the important role of tissue construction and size in determining growth rates and fitness. Here, we show how an expanded optimisation framework, focussed on individual growth rate, enables us to explain shifts in four key traits: leaf mass per area, sapwood area to leaf area ratio (Huber value), wood density and sapwood-specific conductivity in response to soil moisture, atmospheric aridity, CO 2 ${\text{CO}}_{2}$ and light availability. In particular, we predict that as conditions become increasingly dry, height-growth optimising traits shift from resource-acquisitive strategies to resource-conservative strategies, consistent with empirical responses across current environmental gradients of rainfall. These findings can explain both the shift in traits and turnover of species along existing environmental gradients and changing future conditions and highlight the importance of both carbon assimilation and tissue construction in shaping the functional composition of vegetation across climates.

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.

Predicting resilience through the lens of competing adjustments to vegetation function.

There is a pressing need to better understand ecosystem resilience to droughts and heatwaves. Eco-evolutionary optimization approaches have been proposed as means to build this understanding in land surface models and improve their predictive capability, but competing approaches are yet to be tested together. Here, we coupled approaches that optimize canopy gas exchange and leaf nitrogen investment, respectively, extending both approaches to account for hydraulic impairment. We assessed model predictions using observations from a native Eucalyptus woodland that experienced repeated droughts and heatwaves between 2013 and 2020, whilst exposed to an elevated [CO2 ] treatment. Our combined approaches improved predictions of transpiration and enhanced the simulated magnitude of the CO2 fertilization effect on gross primary productivity. The competing approaches also worked consistently along axes of change in soil moisture, leaf area, and [CO2 ]. Despite predictions of a significant percentage loss of hydraulic conductivity due to embolism (PLC) in 2013, 2014, 2016, and 2017 (99th percentile PLC > 45%), simulated hydraulic legacy effects were small and short-lived (2 months). Our analysis suggests that leaf shedding and/or suppressed foliage growth formed a strategy to mitigate drought risk. Accounting for foliage responses to water availability has the potential to improve model predictions of ecosystem resilience.

Integrating the evidence for a terrestrial carbon sink caused by increasing atmospheric CO2.

Atmospheric carbon dioxide concentration ([CO2 ]) is increasing, which increases leaf-scale photosynthesis and intrinsic water-use efficiency. These direct responses have the potential to increase plant growth, vegetation biomass, and soil organic matter; transferring carbon from the atmosphere into terrestrial ecosystems (a carbon sink). A substantial global terrestrial carbon sink would slow the rate of [CO2 ] increase and thus climate change. However, ecosystem CO2 responses are complex or confounded by concurrent changes in multiple agents of global change and evidence for a [CO2 ]-driven terrestrial carbon sink can appear contradictory. Here we synthesize theory and broad, multidisciplinary evidence for the effects of increasing [CO2 ] (iCO2 ) on the global terrestrial carbon sink. Evidence suggests a substantial increase in global photosynthesis since pre-industrial times. Established theory, supported by experiments, indicates that iCO2 is likely responsible for about half of the increase. Global carbon budgeting, atmospheric data, and forest inventories indicate a historical carbon sink, and these apparent iCO2 responses are high in comparison to experiments and predictions from theory. Plant mortality and soil carbon iCO2 responses are highly uncertain. In conclusion, a range of evidence supports a positive terrestrial carbon sink in response to iCO2 , albeit with uncertain magnitude and strong suggestion of a role for additional agents of global change.

Plant profit maximization improves predictions of European forest responses to drought.

Knowledge of how water stress impacts the carbon and water cycles is a key uncertainty in terrestrial biosphere models. We tested a new profit maximization model, where photosynthetic uptake of CO2 is optimally traded against plant hydraulic function, as an alternative to the empirical functions commonly used in models to regulate gas exchange during periods of water stress. We conducted a multi-site evaluation of this model at the ecosystem scale, before and during major droughts in Europe. Additionally, we asked whether the maximum hydraulic conductance in the soil-plant continuum kmax (a key model parameter which is not commonly measured) could be predicted from long-term site climate. Compared with a control model with an empirical soil moisture function, the profit maximization model improved the simulation of evapotranspiration during the growing season, reducing the normalized mean square error by c. 63%, across mesic and xeric sites. We also showed that kmax could be estimated from long-term climate, with improvements in the simulation of evapotranspiration at eight out of the 10 forest sites during drought. Although the generalization of this approach is contingent upon determining kmax , it presents a mechanistic trait-based alternative to regulate canopy gas exchange in global models.

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 assess the vulnerability of Australian trees to drought. Here we implemented a model of plant hydraulics into the Community Atmosphere Biosphere Land Exchange (CABLE) land surface model. We parameterized the drought response behaviour of five broad vegetation types, based on a common garden dry-down experiment with species originating across a rainfall gradient (188-1,125 mm/year) 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) of about 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.

Leaf morphological traits show greater responses to changes in climate than leaf physiological traits and gas exchange variables.

Adaptation to changing conditions is one of the strategies plants may use to survive in the face of climate change. We aimed to determine whether plants' leaf morphological and physiological traits/gas exchange variables have changed in response to recent, anthropogenic climate change. We grew seedlings from resurrected historic seeds from ex-situ seed banks and paired modern seeds in a common-garden experiment. Species pairs were collected from regions that had undergone differing levels of climate change using an emerging framework-Climate Contrast Resurrection Ecology, allowing us to hypothesise that regions with greater changes in climate (including temperature, precipitation, climate variability and climatic extremes) would be greater trait responses in leaf morphology and physiology over time. Our study found that in regions where there were greater changes in climate, there were greater changes in average leaf area, leaf margin complexity, leaf thickness and leaf intrinsic water use efficiency. Changes in leaf roundness, photosynthetic rate, stomatal density and the leaf economic strategy of our species were not correlated with changes in climate. Our results show that leaves do have the ability to respond to changes in climate, however, there are greater inherited responses in morphological leaf traits than in physiological traits/variables and greater responses to extreme measures of climate than gradual changes in climatic means. It is vital for accurate predictions of species' responses to impending climate change to ensure that future climate change ecology studies utilise knowledge about the difference in both leaf trait and gas exchange responses and the climate variables that they respond to.

Time-traveling seeds reveal that plant regeneration and growth traits are responding to climate change.

Studies assessing the biological impacts of climate change typically rely on long-term, historic data to measure trait responses to climate through time. Here, we overcame the problem of absent historical data by using resurrected seeds to capture historic plant-trait data for a number of plant regeneration and growth traits. We collected seed and seedling trait measurements from resurrected historic seeds and compared these with modern seed and seedling traits collected from the same species in the same geographic location. We found a total of 43 species from southeastern Australia for which modern/historic seed pairs could be located. These species were located in a range of regions that have undergone different amounts of climate change across a range of temperature, precipitation, and extreme measures of climate. There was a correlation between the amount of change in climate metrics, and the amount of change in plant traits. Using stepwise model selection, we found that for all regeneration and growth trait changes (except change in stem density), the most accurate model selected at least two measures of climate change. Changes in extreme measures of climate, such as heat-wave duration and changes in climate variability, were more strongly related to changes in regeneration and growth traits than changes in mean climate metrics. Across our species, for every 5% increase in temperature variability, there was a threefold increase in the probability of seed viability and seed germination success. An increase of 1 d in the maximum duration of dry spells through time led to a 1.5-fold decrease in seed viability and seeds became 30% flatter/thinner. Regions where the maximum heat-wave duration had increased by 10 d saw a 1.35-cm decrease in seedling height and a 1.04-g decrease in seedling biomass. Rapid responses in plant traits to changes in climate may be possible; however, it is not clear whether these changes will be fast enough for plants to keep pace with future climate change.