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While tropical cyclone regimes are shifting with climate change, the mechanisms underpinning the resistance (ability to withstand disturbance-induced change) and resilience (capacity to return to pre-disturbance reference) of tropical forest litterfall to cyclones remain largely unexplored pantropically. Single-site studies in Australia and Hawaii suggest that litterfall on low-phosphorus (P) soils is more resistant and less resilient to cyclones. We conducted a meta-analysis to investigate the pantropical importance of total soil P in mediating forest litterfall resistance and resilience to 22 tropical cyclones. We evaluated cyclone-induced and post-cyclone litterfall mass (g/m2 /day), and P and nitrogen (N) fluxes (mg/m2 /day) and concentrations (mg/g), all indicators of ecosystem function and essential for nutrient cycling. Across 73 case studies in Australia, Guadeloupe, Hawaii, Mexico, Puerto Rico, and Taiwan, total litterfall mass flux increased from ~2.5 ± 0.3 to 22.5 ± 3 g/m2 /day due to cyclones, with large variation among studies. Litterfall P and N fluxes post-cyclone represented ~5% and 10% of the average annual fluxes, respectively. Post-cyclone leaf litterfall N and P concentrations were 21.6 ± 1.2% and 58.6 ± 2.3% higher than pre-cyclone means. Mixed-effects models determined that soil P negatively moderated the pantropical litterfall resistance to cyclones, with a 100 mg P/kg increase in soil P corresponding to a 32% to 38% decrease in resistance. Based on 33% of the resistance case studies, total litterfall mass flux reached pre-disturbance levels within one-year post-disturbance. A GAMM indicated that soil P, gale wind duration and time post-cyclone jointly moderate the short-term resilience of total litterfall, with the nature of the relationship between resilience and soil P contingent on time and wind duration. Across pantropical forests observed to date, our results indicate that litterfall resistance and resilience in the face of intensifying cyclones will be partially determined by total soil P.

Lack of tree fecundity data across climatic gradients precludes the analysis of how seed supply contributes to global variation in forest regeneration and biotic interactions responsible for biodiversity. A global synthesis of raw seedproduction data shows a 250-fold increase in seed abundance from cold-dry to warm-wet climates, driven primarily by a 100-fold increase in seed production for a given tree size. The modest (threefold) increase in forest productivity across the same climate gradient cannot explain the magnitudes of these trends. The increase in seeds per tree can arise from adaptive evolution driven by intense species interactions or from the direct effects of a warm, moist climate on tree fecundity. Either way, the massive differences in seed supply ramify through food webs potentially explaining a disproportionate role for species interactions in the wet tropics.

Abstract Droughts are predicted to become more frequent and intense in many tropical regions, which may cause shifts in plant community composition. Especially in diverse tropical communities, understanding how traits mediate demographic responses to drought can help provide insight into the effects of climate change on these ecosystems. To understand tropical tree responses to reduced soil moisture, we grew seedlings of eight species across an experimental soil moisture gradient at the Luquillo Experimental Forest, Puerto Rico. We quantified survival and growth over an 8‐month period and characterized demographic responses in terms of tolerance to low soil moisture—defined as survival and growth rates under low soil moisture conditions—and sensitivity to variation in soil moisture—defined as more pronounced changes in demographic rates across the observed range of soil moisture. We then compared demographic responses with interspecific variation in a suite of 11 (root, stem, and leaf) functional traits, measured on individuals that survived the experiment. Lower soil moisture was associated with reduced survival and growth but traits mediated species‐specific responses. Species with relatively conservative traits (e.g., high leaf mass per area), had higher survival at low soil moisture whereas species with more extensive root systems were more sensitive to soil moisture, in that they exhibited more pronounced changes in growth across the experimental soil moisture gradient. Our results suggest that increasing drought will favor species with more conservative traits that confer greater survival in low soil moisture conditions.

The growth and survival of individual trees determine the physical structure of a forest with important consequences for forest function. However, given the diversity of tree species and forest biomes, quantifying the multitude of demographic strategies within and across forests and the way that they translate into forest structure and function remains a significant challenge. Here, we quantify the demographic rates of 1961 tree species from temperate and tropical forests and evaluate how demographic diversity (DD) and demographic composition (DC) differ across forests, and how these differences in demography relate to species richness, aboveground biomass (AGB), and carbon residence time. We find wide variation in DD and DC across forest plots, patterns that are not explained by species richness or climate variables alone. There is no evidence that DD has an effect on either AGB or carbon residence time. Rather, the DC of forests, specifically the relative abundance of large statured species, predicted both biomass and carbon residence time. Our results demonstrate the distinct DCs of globally distributed forests, reflecting biogeography, recent history, and current plot conditions. Linking the DC of forests to resilience or vulnerability to climate change, will improve the precision and accuracy of predictions of future forest composition, structure, and function.

Abstract One mechanism proposed to explain high species diversity in tropical systems is strong negative conspecific density dependence (CDD), which reduces recruitment of juveniles in proximity to conspecific adult plants. Although evidence shows that plant-specific soil pathogens can drive negative CDD, trees also form key mutualisms with mycorrhizal fungi, which may counteract these effects. Across 43 large-scale forest plots worldwide, we tested whether ectomycorrhizal tree species exhibit weaker negative CDD than arbuscular mycorrhizal tree species. We further tested for conmycorrhizal density dependence (CMDD) to test for benefit from shared mutualists. We found that the strength of CDD varies systematically with mycorrhizal type, with ectomycorrhizal tree species exhibiting higher sapling densities with increasing adult densities than arbuscular mycorrhizal tree species. Moreover, we found evidence of positive CMDD for tree species of both mycorrhizal types. Collectively, these findings indicate that mycorrhizal interactions likely play a foundational role in global forest diversity patterns and structure.

The relationships that control seed production in trees are fundamental to understanding the evolution of forest species and their capacity to recover from increasing losses to drought, fire, and harvest. A synthesis of fecundity data from 714 species worldwide allowed us to examine hypotheses that are central to quantifying reproduction, a foundation for assessing fitness in forest trees. Four major findings emerged. First, seed production is not constrained by a strict trade-off between seed size and numbers. Instead, seed numbers vary over ten orders of magnitude, with species that invest in large seeds producing more seeds than expected from the 1:1 trade-off. Second, gymnosperms have lower seed production than angiosperms, potentially due to their extra investments in protective woody cones. Third, nutrient-demanding species, indicated by high foliar phosphorus concentrations, have low seed production. Finally, sensitivity of individual species to soil fertility varies widely, limiting the response of community seed production to fertility gradients. In combination, these findings can inform models of forest response that need to incorporate reproductive potential.

Despite its importance for forest regeneration, food webs, and human economies, changes in tree fecundity with tree size and age remain largely unknown. The allometric increase with tree diameter assumed in ecological models would substantially overestimate seed contributions from large trees if fecundity eventually declines with size. Current estimates are dominated by overrepresentation of small trees in regression models. We combined global fecundity data, including a substantial representation of large trees. We compared size-fecundity relationships against traditional allometric scaling with diameter and two models based on crown architecture. All allometric models fail to describe the declining rate of increase in fecundity with diameter found for 80% of 597 species in our analysis. The strong evidence of declining fecundity, beyond what can be explained by crown architectural change, is consistent with physiological decline. A downward revision of projected fecundity of large trees can improve the next generation of forest dynamic models.

Arbuscular mycorrhizal (AM) and ectomycorrhizal (EcM) associations are critical for host-tree performance. However, how mycorrhizal associations correlate with the latitudinal tree beta-diversity remains untested. Using a global dataset of 45 forest plots representing 2,804,270 trees across 3840 species, we test how AM and EcM trees contribute to total beta-diversity and its components (turnover and nestedness) of all trees. We find AM rather than EcM trees predominantly contribute to decreasing total beta-diversity and turnover and increasing nestedness with increasing latitude, probably because wide distributions of EcM trees do not generate strong compositional differences among localities. Environmental variables, especially temperature and precipitation, are strongly correlated with beta-diversity patterns for both AM trees and all trees rather than EcM trees. Results support our hypotheses that latitudinal beta-diversity patterns and environmental effects on these patterns are highly dependent on mycorrhizal types. Our findings highlight the importance of AM-dominated forests for conserving global forest biodiversity.

Abstract Projected increases in hurricane intensity under a warming climate will have profound effects on many forest ecosystems. One key challenge is to disentangle the effects of wind damage from the myriad factors that influence forest structure and species distributions over large spatial scales. Here, we employ a novel machine learning framework with high‐resolution aerial photos, and LiDAR collected over 115 km2 of El Yunque National Forest in Puerto Rico to examine the effects of topographic exposure to two hurricanes, Hugo (1989) and Georges (1998), and several landscape‐scale environmental factors on the current forest height and abundance of a dominant, wind‐resistant species, the palm Prestoea acuminata var. montana. Model predictions show that the average density of the palm was 32% greater while the canopy height was 20% shorter in forests exposed to the two storms relative to unexposed areas. Our results demonstrate that hurricanes have lasting effects on forest canopy height and composition, suggesting the expected increase in hurricane severity with a warming climate will alter coastal forests in the North Atlantic.

Abstract Collaboration between ecologists and learning scientists can give rise to powerful models for scientific outreach within ecology. This paper presents a process by which learning scientists and ecologists codesigned a science curriculum that invites students to join an ecological community of practice. In the Journey to El Yunque middle school science curriculum, students engage with simulation models generated from data gathered by Luquillo Long Term Ecological Research (LUQ LTER) scientists. Journey to El Yunque students can explore post‐hurricane population changes in yagrumo (Cecropia schreberiana), tabonuco (Dacryodes excelsa), coquís (Eleutherodactylus coquí), snails (Caracolus caracola), anoles (Anolis stratulus and A. gundlachi), veiled stinkhorn mushrooms (Dictyophora indusiata), and caterpillars (Historis odius). Ecology‐based revisions to Journey to El Yunque have included adding models of the effects of repeated hurricanes on limiting factors, based in part on findings from a canopy trimming experiment. Revisions based on classroom testing include simplifying student‐facing model controls to allow students to focus on the essential model components. The ongoing collaboration that keeps the Journey to El Yunque curriculum on the cutting edge of ecological and educational advances has been sustained for over two decades. We attribute the longevity of this work to (1) the long‐term nature of LUQ LTER, (2) a sustained interdisciplinary collaboration, and (3) our long‐term relationships with schools.

The relationships that control seed production in trees are key to understand evolutionary pressures that have shaped forests. A global synthesis of fecundity data reveals that while seed production is not constrained by a strict size-number trade-off, it is influenced by taxonomy and nutrient allocation.

Adaptively managing marine protected areas (MPAs) requires accurately assessing whether established MPAs are achieving their goals of protecting and conserving biomass, especially for harvested populations. Ecological MPA assessments commonly compare inside of the MPA to a reference point outside of and/or before implementation (i.e., calculating "response ratios"). Yet, MPAs are not simple ecological experiments; by design, protected populations interact with those outside, and population dynamic responses can be nonlinear. This complicates assessment interpretations. Here, we used a two‐patch population model to explore how MPA response ratios (outside–inside, before–after, and before–after‐control‐impact [BACI]) for fished populations behave under different conditions, like whether the population is receiving a sustainable larval supply or if it is declining despite protection from harvest. We then conducted a Bayesian evaluation of MPA effects on fish and invertebrate populations based on data collected from 82 published studies on 264 no‐take MPAs worldwide, using the results of an earlier global meta‐analysis as priors. We considered the effects of calculating different summary metrics on these results, drawing on the theoretical insights from our population model as a comparative framework. We demonstrate that not all response ratio comparison types provide the same information: For example, outside–inside and BACI comparisons can fail to detect population decline within MPAs, whereas before–after comparisons likely detect that pattern. Considering these limitations, we nonetheless found that MPAs globally are producing positive outcomes, with on average greater biomass, density, and organism size within their boundaries than reference sites. However, only a small portion of studies (18 of 82) provided the temporal data necessary to determine that protection, on average, has led to increased abundance of populations within MPAs over time. These findings demonstrate the importance of considering the underlying system dynamics when assessing MPA effects. Assuming that large outside–inside or BACI response ratios always reflect large and net positive conservation effects may lead to misleading conclusions, we recommend that: (1) when assessing specific MPA effects, empirical findings be considered alongside theoretical knowledge relevant to that MPA system, and (2) management should respond to the local conditions and outcomes, rather than a blanket expectation for positive MPA effects. [ABSTRACT FROM AUTHOR]

As the developers of "Journey to El Yunque," we have taken a different approach to the process of designing a science curriculum. Rather than start with a specific set of concepts or skills to target as learning outcomes, we started by identifying a specific community of practice to which we sought to connect students. Researchers in the El Yunque rainforest in Puerto Rico have been studying the impact of hurricanes on ecosystem dynamics and have been modeling what the long-term impact would be if changes to the global climate increase the frequency of severe hurricanes. Therefore, hurricane impact became the focal phenomenon for the unit. We modeled the process of investigating hurricane impact after the long-term ecological research practices of researchers in El Yunque. Students begin by investigating the long-term impact of hurricanes on the producers in El Yunque. Next students investigate the long-term impact of hurricanes on various consumers in the rainforest. Finally, students investigate how hurricanes impact the cycling of resources directly as well as indirectly through changes in organisms' use of those resources in the rainforest. A central tension in the design process is how to coherently represent the spatial relationships between the components of the ecosystem and the temporal dynamics of the individual components. In this paper, we present the evolution of the program as we sought to balance that design tension and build an environment that connects students to the central phenomenon and practices of the community of researchers in El Yunque.

The nutrient demands of regrowing tropical forests are partly satisfied by nitrogen-fixing legume trees, but our understanding of the abundance of those species is biased towards wet tropical regions. Here we show how the abundance of Leguminosae is affected by both recovery from disturbance and large-scale rainfall gradients through a synthesis of forest inventory plots from a network of 42 Neotropical forest chronosequences. During the first three decades of natural forest regeneration, legume basal area is twice as high in dry compared with wet secondary forests. The tremendous ecological success of legumes in recently disturbed, water-limited forests is likely to be related to both their reduced leaflet size and ability to fix N2, which together enhance legume drought tolerance and water-use efficiency. Earth system models should incorporate these large-scale successional and climatic patterns of legume dominance to provide more accurate estimates of the maximum potential for natural nitrogen fixation across tropical forests.

Abstract Studies of spatial point patterns (SPPs) are often used to examine the role that density‐dependence (DD) and environmental filtering (EF) play in community assembly and species coexistence in forest communities. However, SPP analyses often struggle to distinguish the opposing effects that DD and EF may have on the distribution of tree species. We tested percolation threshold analysis on simulated tree communities as a method to distinguish the importance of thinning from DD EF on SPPs. We then compared the performance of percolation threshold analysis results and a Gibbs point process model in detecting environmental associations as well as clustering patterns or overdispersion. Finally, we applied percolation threshold analysis and the Gibbs point process model to observed SPPs of 12 dominant tree species in a Puerto Rican forest to detect evidence of DD and EF. Percolation threshold analysis using simulated SPPs detected a decrease in clustering due to DD and an increase in clustering from EF. In contrast, the Gibbs point process model clearly detected the effects of EF but only identified DD thinning in two of the four types of simulated SPPs. Percolation threshold analysis on the 12 observed tree species' SPPs found that the SPPs for two species were consistent with thinning from DD processes only, four species had SPPs consistent with EF only and SPP for five reflected a combination of both processes. Gibbs models of observed SPPs of living trees detected significant environmental associations for 11 species and clustering consistent with DD processes for seven species. Percolation threshold analysis is a robust method for detecting community assembly processes in simulated SPPs. By applying percolation threshold analysis to natural communities, we found that tree SPPs were consistent with thinning from both DD and EF. Percolation threshold analysis was better suited to detect DD thinning than Gibbs models for clustered simulated communities. Percolation threshold analysis improves our understanding of forest community assembly processes by quantifying the relative importance of DD and EF in forest communities.

Abstract The ‘Sepsis Six’ bundle was promoted as a deliverable tool outside of the critical care settings, but there is very little data available on the progress and change of sepsis care outside the critical care environment in the UK. Our aim was to compare the yearly prevalence, outcome and the Sepsis Six bundle compliance in patients at risk of mortality from sepsis in non-intensive care environments. Patients with a National Early Warning Score (NEWS) of 3 or above and suspected or proven infection were enrolled into four yearly 24-h point prevalence studies, carried out in fourteen hospitals across Wales from 2016 to 2019. We followed up patients to 30 days between 2016–2019 and to 90 days between 2017 and 2019. Out of the 26,947 patients screened 1651 fulfilled inclusion criteria and were recruited. The full ‘Sepsis Six’ care bundle was completed on 223 (14.0%) occasions, with no significant difference between the years. On 190 (11.5%) occasions none of the bundle elements were completed. There was no significant correlation between bundle element compliance, NEWS or year of study. One hundred and seventy (10.7%) patients were seen by critical care outreach; the ‘Sepsis Six’ bundle was completed significantly more often in this group (54/170, 32.0%) than for patients who were not reviewed by critical care outreach (168/1385, 11.6%; p

Abstract Tropical forest understory regeneration occurs rapidly after disturbance with compositional trajectories that depend on species availability and environmental conditions. To predict future tropical forest regeneration dynamics, we need a deeper understanding of how pulse disturbance events, like hurricanes, interact with environmental variability to affect understory demography and composition. We examined fern and sapling mortality, recruitment, and community composition in relation to solar radiation and soil moisture using 17 years of forest dynamics data (2003–2019) from the Canopy Trimming Experiment in the Luquillo Experimental Forest, Puerto Rico. Solar radiation increased 150% and soil moisture increased 40% following canopy trimming of experimental plots relative to control plots. All plots were disturbed in 2017 by Hurricanes Irma and Maria, so experimentally trimmed plots presented the opportunity to study the effects of multiple hurricanes, while control plots isolated the effects of a single natural hurricane. Recruitment rates maximized at 0.14 individuals/plot/month for ferns and 0.20 stems/plot/month for saplings. Recruitment and mortality were distributed more evenly over the 17 years of monitoring in experimentally trimmed plots than in control plots; however, following Hurricane Maria demographic rates substantially increased in control plots only. In experimentally trimmed plots, the largest community compositional shifts occurred as a result of the trimming events, and compositional changes were greatest for control plots after Hurricane Maria in 2017. Pioneer tree and fern species increased in abundance in response to both simulated and natural hurricanes. Following Hurricane Maria, two dominant pioneer species, Cyathea arborea and Cecropia schreberiana, recruited abundantly, but only in control plots. In trimmed plots, increased solar radiation and soil moisture shifted understory species composition steadily toward pioneer and secondary‐successional species, with soil moisture interacting strongly with canopy trimming. Thus, both solar radiation and soil moisture are environmental drivers affecting pioneer species recruitment following disturbance, which interact with canopy opening following hurricanes. Our results suggest that if hurricane disturbances increase in frequency and severity, as suggested by climate change predictions, the understory regeneration of late‐successional species, such as Manilkara bidentata and Sloanea berteroana, which prefer deeper shade and slightly drier soil microsites, may become imperiled.

The relationship of mycorrhizal associations with latitudinal gradients in tree beta-diversity is unexplored. Using a global dataset approach, this study examines how trees with arbuscular mycorrhizal and ectomycorrhizal associations contribute to latitudinal beta-diversity patterns and the environmental controls of these patterns.

Abstract Climate change and disturbance make it difficult to project long‐term patterns of carbon sequestration in tropical forests, but large ecosystem experiments in these forests can inform predictions. The Canopy Trimming Experiment (CTE) manipulates two key components of hurricane disturbance, canopy openness and detritus deposition, in a tropical forest in Puerto Rico. We documented how the CTE and a real hurricane affected tree recruitment, biomass, and aboveground carbon storage over 15 years. In the CTE treatments, we trimmed branches, but we did not fell trees. We expected that during the 14‐year period after initial canopy trimming, regrowth of branches and stems and stem recruitment stimulated by increased light and trimmed debris would help restore biomass and carbon loss due to trimming. Compared to control plots, in the trimmed plots recruitment of palms and dicot trees increased markedly after trimming, and stem diameters of standing trees increased. Data showed that recruitment of small trees adds little to aboveground carbon, compared to the amount in large trees. Nevertheless, this response restored pretreatment biomass and carbon in the experimental period. In particular, the experimental additions of trimmed debris on the forest floor seemed to stimulate increase in aboveground carbon. Toward the end of the experimental period, Hurricane Maria (Category 4 hurricane) trimmed and felled some trees but reduced aboveground carbon less in the plots (including untrimmed plots) than experimental trimming had. Thus, it appears that the amount of regrowth recorded after experimental trimming could also restore aboveground carbon in the forest after a severe hurricane in the same time span. However, Hurricane Maria, unlike the trimming treatments, felled large trees, and it may be that with predicted, more frequent severe hurricanes, that the continued loss of large trees would over the long term decrease aboveground carbon stored in this Puerto Rican forest and likewise in other tropical forests affected by cyclonic storms.

The ability to predict growth and potential yield is essential for planning forest management. Here we developed a narrow size-class diameter model to simulate growth in a mature plantation (age 42 years) of big-leaf mahogany (Swietenia macrophylla King) in a subtropical moist forest in Puerto Rico. We calculated optimal rotation age by means of the biological and economic criteria. The analyses involved diameter measurements over 31 years and a variety of contrasting economic scenarios, which were evaluated by means of the Faustman formula and sensitivity analyses. The optimal biological rotation was 90 years while the optimal economic rotation was 42 years in most of the scenarios, regardless of wood prices and costs of reforestation. Conspicuous changes in the optimal economic rotation were only observed under low interest rates, in which it was prolonged to around 80 years, close to the optimal biological rotation. Abundant natural regeneration and a prolonged biological rotation suggest that other silvicultural systems, rather than traditional clear-cutting and replanting could be adopted. However, to evaluate alternative silvicultural strategies we would need to improve our ability to predict growth. This would require expansion of our database to include information on growth conditions of individual trees and young tree dynamics, as well as size and growth of other species, to model natural regeneration and competition.

Given the potential for increasingly common and intense tropical storms, it is important to understand their effects on island forest communities. Here, the authors show that Hurricane María’s strength and rainfall had larger effects on tree mortality than other less severe storms, and that large trees and species with low-density wood were most susceptible.

Native flora and fauna of Puerto Rico have a long biogeographic connection to South America. Theory and empirical evidence suggest that islands, particularly those distantly isolated from the mainland, should be more susceptible to naturalizations and invasions of non-native species than continental areas. Anthropogenic disturbances can facilitate accidental and deliberate introductions of non-native species. In this study, we asked: What is the current status of introduced species within El Yunque National Forest (EYNF), the largest and most well-conserved forest area of Puerto Rico? To address this question, we reviewed the literature and surveyed local experts to identify introduced plant and animal taxa that are behaving as invaders within EYNF. We hypothesized that well-conserved forest areas within EYNF would be more resistant to invasions than disturbed areas along roads and ruderal areas with a long history of human activity. We found that there is only partial evidence that supports our hypothesis and this evidence is strongest in vascular plants, but not for the other taxonomic groups analyzed. Our combined results showed that currently the more ubiquitous invasive species in EYNF include some mammals (feral cat, rat, and mongoose) and some invertebrates (earthworms, mosquito, and Africanized honeybee). For many taxa, there is little information to thoroughly test our hypothesis, and thus more detailed surveys of the status of non-native and invasive species in EYNF are needed.

Predicting drought responses of individual trees in tropical forests remains challenging, in part because trees experience drought differently depending on their position in spatially heterogeneous environments. Specifically, topography and the competitive environment can influence the severity of water stress experienced by individual trees, leading to individual-level variation in drought impacts. A drought in 2015 in Puerto Rico provided the opportunity to assess how drought response varies with topography and neighborhood crowding in a tropical forest. In this study, we integrated 3 years of annual census data from the El Yunque Chronosequence plots with measurements of functional traits and LiDAR-derived metrics of microsite topography. We fit hierarchical Bayesian models to examine how drought, microtopography, and neighborhood crowding influence individual tree growth and survival, and the role functional traits play in mediating species’ responses to these drivers. We found that while growth was lower during the drought year, drought had no effect on survival, suggesting that these forests are fairly resilient to a single-year drought. However, growth response to drought, as well as average growth and survival, varied with topography: tree growth in valley-like microsites was more negatively affected by drought, and survival was lower on steeper slopes while growth was higher in valleys. Neighborhood crowding reduced growth and increased survival, but these effects did not vary between drought/non-drought years. Functional traits provided some insight into mechanisms by which drought and topography affected growth and survival. For example, trees with high specific leaf area grew more slowly on steeper slopes, and high wood density trees were less sensitive to drought. However, the relationships between functional traits and response to drought and topography were weak overall. Species sorting across microtopography may drive observed relationships between average performance, drought response, and topography. Our results suggest that understanding species’ responses to drought requires consideration of the microenvironments in which they grow. Complex interactions between regional climate, topography, and traits underlie individual and species variation in drought response.