Numerous current efforts seek to improve the representation of ecosystem ecology and vegetation demographic processes within Earth System Models (ESMs). These developments are widely viewed as an important step in developing greater realism in predictions of future ecosystem states and fluxes. Increased realism, however, leads to increased model complexity, with new features raising a suite of ecological questions that require empirical constraints. Here, we review the developments that

We assessed whether diversity in plant hydraulic traits can explain the observed diversity in plant responses to water stress in seasonally dry tropical forests (SDTFs). The Ecosystem Demography model 2 (ED2) was updated with a trait-driven mechanistic plant hydraulic module, as well as novel drought-phenology and plant water stress schemes. Four plant functional types were parameterized on the basis of meta-analysis of plant hydraulic traits. Simulations from both the original and the updated ED2 were evaluated against 5 yr of field data from a Costa Rican SDTF site and remote-sensing data over Central America. The updated model generated realistic plant hydraulic dynamics, such as leaf water potential and stem sap flow. Compared with the original ED2, predictions from our novel trait-driven model matched better with observed growth, phenology and their variations among functional groups. Most notably, the original ED2 produced unrealistically small leaf area index (LAI) and underestimated cumulative leaf litter. Both of these biases were corrected by the updated model. The updated model was also better able to simulate spatial patterns of LAI dynamics in Central America. Plant hydraulic traits are intercorrelated in SDTFs. Mechanistic incorporation of plant hydraulic traits is necessary for the simulation of spatiotemporal patterns of vegetation dynamics in SDTFs in vegetation models.

Summary Tree mortality rates appear to be increasing in moist tropical forests (MTFs) with significant carbon cycle consequences. Here, we review the state of knowledge regarding MTF tree mortality, create a conceptual framework with testable hypotheses regarding the drivers, mechanisms and interactions that may underlie increasing MTF mortality rates, and identify the next steps for improved understanding and reduced prediction. Increasing mortality rates are associated with rising temperature and vapor pressure deficit, liana abundance, drought, wind events, fire and, possibly, CO2 fertilization‐induced increases in stand thinning or acceleration of trees reaching larger, more vulnerable heights. The majority of these mortality drivers may kill trees in part through carbon starvation and hydraulic failure. The relative importance of each driver is unknown. High species diversity may buffer MTFs against large‐scale mortality events, but recent and expected trends in mortality drivers give reason for concern regarding increasing mortality within MTFs. Models of tropical tree mortality are advancing the representation of hydraulics, carbon and demography, but require more empirical knowledge regarding the most common drivers and their subsequent mechanisms. We outline critical datasets and model developments required to test hypotheses regarding the underlying causes of increasing MTF mortality rates, and improve prediction of future mortality under climate change.

Seasonally dry tropical forests (SDTF) are located in regions with alternating wet and dry seasons, with dry seasons that last several months or more. By the end of the 21st century, climate models predict substantial changes in rainfall regimes across these regions, but little is known about how individuals, species, and communities in SDTF will cope with the hotter, drier conditions predicted by climate models. In this review, we explore different rainfall scenarios that may result in ecological drought in SDTF through the lens of two alternative hypotheses: 1) these forests will be sensitive to drought because they are already limited by water and close to climatic thresholds, or 2) they will be resistant/resilient to intra-and inter-annual changes in rainfall because they are adapted to predictable, seasonal drought. In our review of literature that spans microbial to ecosystem processes, a majority of the available studies suggests that increasing frequency and intensity of droughts in SDTF will likely alter species distributions and ecosystem processes. Though we conclude that SDTF will be sensitive to altered rainfall regimes, many gaps in the literature remain. Future research should focus on geographically comparative studies and well-replicated drought experiments that can provide empirical evidence to improve simulation models used to forecast SDTF responses to future climate change at coarser spatial and temporal scales.

Drought-related tree mortality is now a widespread phenomenon predicted to increase in magnitude with climate change. However, the patterns of which species and trees are most vulnerable to drought, and the underlying mechanisms have remained elusive, in part due to the lack of relevant data and difficulty of predicting the location of catastrophic drought years in advance. We used long-term demographic records and extensive databases of functional traits and distribution patterns to understand the responses of 20-53 species to an extreme drought in a seasonally dry tropical forest in Costa Rica, which occurred during the 2015 El Niño Southern Oscillation event. Overall, species-specific mortality rates during the drought ranged from 0% to 34%, and varied little as a function of tree size. By contrast, hydraulic safety margins correlated well with probability of mortality among species, while morphological or leaf economics spectrum traits did not. This firmly suggests hydraulic traits as targets for future research. K E Y W O R D S extreme drought, hydraulic traits, rainfall seasonality, tree mortality | 3123 POWERS Et al.

This paper presents a continental-scale phenological analysis of African savannas and woodlands.We apply an array of synergistic vegetation and hydrological data records from satellite remote sensing and model simulations to explore the influence of rainy season timing and duration on regional land surface phenology and ecosystem structure. We find that (i) the rainy season onset precedes and is an effective predictor of the growing season onset in African grasslands. (ii) African woodlands generally have early green-up before rainy season onset and have a variable delayed senescence period after the rainy season, with this delay correlated nonlinearly with tree fraction. These woodland responses suggest their complex water use mechanisms (either from potential groundwater use by relatively deep roots or stem-water reserve) to maintain dry season activity. (iii) We empirically find that the rainy season length has strong nonlinear impacts on tree fractional cover in the annual rainfall range from 600 to 1800 mm/yr, which may lend some support to the previous modeling study that given the same amount of total rainfall to the tree fraction may first increase with the lengthening of rainy season until reaching an "optimal rainy season length," after which tree fraction decreases with the further lengthening of rainy season. This nonlinear response is resulted from compound mechanisms of hydrological cycle, fire, and other factors. We conclude that African savannas and deciduous woodlands have distinctive responses in their phenology and ecosystem functioning to rainy season. Further research is needed to address interaction between groundwater and tropical woodland as well as to explicitly consider the ecological significance of rainy season length under climate change.

Gross ecosystem productivity (GEP) in tropical forests varies both with the environment and with biotic changes in photosynthetic infrastructure, but our understanding of the relative effects of these factors across timescales is limited. Here, we used a statistical model to partition the variability of seven years of eddy covariance-derived GEP in a central Amazon evergreen forest into two main causes: variation in environmental drivers (solar radiation, diffuse light fraction, and vapor pressure deficit) that interact with model parameters that govern photosynthesis and biotic variation in canopy photosynthetic light-use efficiency associated with changes in the parameters themselves. Our fitted model was able to explain most of the variability in GEP at hourly (R = 0.77) to interannual (R = 0.80) timescales. At hourly timescales, we found that 75% of observed GEP variability could be attributed to environmental variability. When aggregating GEP to the longer timescales (daily, monthly, and yearly), however, environmental variation explained progressively less GEP variability: At monthly timescales, it explained only 3%, much less than biotic variation in canopy photosynthetic light-use efficiency, which accounted for 63%. These results challenge modeling approaches that assume GEP is primarily controlled by the environment at both short and long timescales. Our approach distinguishing biotic from environmental variability can help to resolve debates about environmental limitations to tropical forest photosynthesis. For example, we found that biotically regulated canopy photosynthetic light-use efficiency (associated with leaf phenology) increased with sunlight during dry seasons (consistent with light but not water limitation of canopy development) but that realized GEP was nonetheless lower relative to its potential efficiency during dry than wet seasons (consistent with water limitation of photosynthesis in given assemblages of leaves). This work highlights the importance of accounting for differential regulation of GEP at different timescales and of identifying the underlying feedbacks and adaptive mechanisms.

Tree abundance in tropical savannas exhibits large and unexplained spatial variability. Here, we propose that differentiated tree and grass water use strategies can explain the observed negative relation between maximum tree abundance and rainfall intensity (defined as the characteristic rainfall depth on rainy days), and we present a biophysical tree-grass competition model to test this idea. The model is founded on a premise that has been well established in empirical studies, namely, that the relative growth rate of grasses is much higher compared with trees in wet conditions but that grasses are more susceptible to water stress and lose biomass more quickly in dry conditions. The model is coupled with a stochastic rainfall generator and then calibrated and tested using field observations from several African savanna sites. We show that the observed negative relation between maximum tree abundance and rainfall intensity can be explained only when differentiated water use strategies are accounted for. Numerical experiments reveal that this effect is more significant than the effect of root niche separation. Our results emphasize the importance of vegetation physiology in determining the responses of tree abundance to climate variations in tropical savannas and suggest that projected increases in rainfall intensity may lead to an increase in grass in this biome.savanna | tree abundance | rainfall intensity | plant water use | competition D espite the important role of disturbance (1-3), climatic variables, especially precipitation, are considered to be the primary determinants of the maximum achievable tree abundance in tropical savannas (4,5). Most empirical studies consider the effects of mean annual precipitation (MAP) (4-6) but not precipitation variability. However, evidence is emerging that precipitation variability is also critically important for determining savanna tree abundance (7-9). In a satellite-based study that included all African savannas, maximum tree cover was found to decrease with increasing rainfall intensity after MAP was controlled for (7). This pattern points to a fundamental, but as yet unarticulated, mechanistic relation between rainfall intensity and tree abundance. This result also seems incompatible with classical root niche separation theory (10, 11). The theory posits that trees have deeper roots than grasses. Trees would have a relative advantage over grasses under more intense rainfall scenario because more water infiltrates into deep soil layer. Therefore, maximum tree cover should increase with increasing rainfall intensity if the classical root niche separation theory holds true.In dry savannas (MAP < 1,000 mm), daily rainfall variability leads to pronounced stochasticity in the amount of soil water available to plants. The capability of efficiently using water during wet periods and while also surviving dry periods (here referred to as a water use strategy) largely defines plant competitiveness and therefore plant abundance. Here, we parameterize a biophysical tree-grass c...