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Drought and nutrient input are two main global change drivers that threaten ecosystem function and services. Resolving the interactive effects of human-induced stressors on individual species is necessary to improve our understanding of community and ecosystem responses. This study comparatively assessed how different nutrient conditions affect whole-plant drought responses across 13 common temperate grassland species. We conducted a fully factorial drought-fertilization experiment to examine the effect of nutrient addition [nitrogen (N), phosphorus (P), and combined NP] on species' drought survival, and on drought resistance of growth as well as drought legacy effects. Drought had an overall negative effect on survival and growth, and the adverse drought effects extended into the next growing season. Neither drought resistance nor legacy effects exhibited an overall effect of nutrients. Instead, both the size and the direction of the effects differed strongly among species and between nutrient conditions. Consistently, species performance ranking under drought changed with nitrogen availability. The idiosyncratic responses of species to drought under different nutrient conditions may underlie the seemingly contradicting effects of drought in studies on grassland composition and productivity along nutrient and land-use gradients—ranging from amplifying to dampening. Differential species’ responses to combinations of nutrients and drought, as observed in our study, complicate predictions of community and ecosystem responses to climate and land-use changes. Moreover, they highlight the urgent need for an improved understanding of the mechanisms that render species more or less vulnerable to drought under different nutrients.

As extreme climate events are predicted to become more frequent because of global climate change, understanding their impacts on natural systems is crucial. Tropical forests are vulnerable to droughts associated with extreme El Niño events. However, little is known about how tropical seedling communities respond to El Niño-related droughts, even though patterns of seedling survival shape future forest structure and diversity. Using long-term data from eight tropical moist forests spanning a rainfall gradient in central Panama, we show that community-wide seedling mortality increased by 11% during the extreme 2015-16 El Niño, with mortality increasing most in drought-sensitive species and in wetter forests. These results indicate that severe El Niño-related droughts influence understory dynamics in tropical forests, with effects varying both within and across sites. Our findings suggest that predicted increases in the frequency of extreme El Niño events will alter tropical plant communities through their effects on early life stages.

Silicon (Si) accumulation is known to alleviate various biotic and abiotic stressors in plants with potential ecological consequences. However, for dicotyledonous plants our understanding of Si variation remains limited. We conducted a comparative experimental study to investigate (1) interspecific variation of foliar Si concentrations across 37 dicotyledonous forbs of temperate grasslands, (2) intraspecific variation in foliar Si concentration in response to soil Si availability, the influence of (3) phylogenetic relatedness, and (4) habitat association to moisture. Foliar Si differed markedly (approx. 70-fold) across the investigated forbs, with some species exhibiting Si accumulation similar to grasses. Foliar Si increased with soil Si availability, but the response varied across species: species with higher Si accumulation capacity showed a stronger response, indicating that they did not actively upregulate Si uptake under low soil Si availability. Foliar Si showed a pronounced phylogenetic signal, i.e., closely related species exhibited more similar foliar Si concentrations than distantly related species. Significant differences in foliar Si concentration within closely related species pairs nevertheless support that active Si uptake and associated high Si concentrations has evolved multiple times in forbs. Foliar Si was not higher in species associated with drier habitats, implying that in dicotyledonous forbs of temperate grasslands high foliar Si is not an adaptive trait to withstand drought. Our results demonstrated considerable inter- and intraspecific variation in foliar Si concentration in temperate forbs. This variation should have pervasive, but so far understudied, ecological consequences for community composition and functioning of temperate grasslands under land-use and climate change. Supplementary Information The online version contains supplementary material available at 10.1007/s00442-021-04978-9.

Aim: Here we examine the functional profile of regional tree species pools across the latitudinal distribution of Neotropical moist forests, and test trait–climate relationships among local communities. We expected opportunistic strategies (acquisitive traits, small seeds) to be overrepresented in species pools further from the equator, but also in terms of abundance in local communities in currently wetter, warmer and more seasonal climates. Location: Neotropics. Time period: Recent. Major taxa studied: Trees. Methods: We obtained abundance data from 471 plots across nine Neotropical regions, including c. 100,000 trees of 3,417 species, in addition to six functional traits. We compared occurrence-based trait distributions among regional species pools, and evaluated single trait–climate relationships across local communities using community abundance-weighted means (CWMs). Multivariate trait–climate relationships were assessed by a double-constrained correspondence analysis that tests both how CWMs relate to climate and how species distributions, parameterized by niche centroids in climate space, relate to their traits. Results: Regional species pools were undistinguished in functional terms, but opportunistic strategies dominated local communities further from the equator, particularly in the Northern Hemisphere. Climate explained up to 57% of the variation in CWM traits, with increasing prevalence of lower-statured, light-wooded and softer-leaved species bearing smaller seeds in more seasonal, wetter and warmer climates. Species distributions were significantly but weakly related to functional traits. Main conclusions: Neotropical moist forest regions share similar sets of functional strategies, from which local assembly processes, driven by current climatic conditions, select for species with different functional strategies. We can thus expect functional responses to climate change driven by changes in relative abundances of species already present regionally. Particularly, equatorial forests holding the most conservative traits and large seeds are likely to experience the most severe changes if climate change triggers the proliferation of opportunistic tree species.

Identifying key traits that can serve as proxies for species drought resistance is crucial for predicting and mitigating the effects of climate change in diverse plant communities. Turgor loss point (π

Turgor loss point (πₜₗₚ) has been suggested to be a key trait for drought resistance in woody species. In herbaceous grassland species, the role of πₜₗₚ for species drought survival has not yet been tested, although grasslands are projected to experience more frequent and intense droughts with climate change. To gain insights into the role of πₜₗₚ for drought resistance of temperate perennial grassland species, we assessed πₜₗₚ of 41 species common in Germany (20 forbs, 21 grasses). We directly related them to the species’ comparative whole‐plant drought survival and midday leaf water potentials under drought (ΨMD) assessed in a common garden drought experiment, and to species moisture association. Species drought survival increased with increasing πₜₗₚ across all species as well as within forbs or grasses separately. ΨMD was positively related to πₜₗₚ and drought survival. Our results imply that high πₜₗₚ promotes drought survival of common perennial European temperate mesic grassland species by enabling them to maintain high leaf water potentials under drought, that is, a desiccation avoidance strategy. However, πₜₗₚ was not related to species moisture association. The positive relationship between πₜₗₚ and drought survival in herbaceous grassland species was opposite to the negative relationship previously established in woody plants, implying that mechanisms of drought resistance differ between woody and herbaceous species. Our results highlight the necessity of directly testing the relationship of functional traits to whole‐plant drought survival in different plant life forms, before using trait assessments for predicting plant responses to drought. A free plain language summary can be found within the Supporting Information of this article.

To improve projections of consequences of increasing intensity and frequency of drought events for grasslands, we need a thorough understanding of species performance responses to drought, of performance trade-offs and how drought resistance is related to species distributions. However, comparative and quantitative assessments of whole-plant drought resistance that allow to rigorously address these aspects are lacking for temperate grassland species. We conducted a common garden experiment with 40 common temperate grassland species to compare species survival and growth under intense drought and well-irrigated conditions. Overall, survival and growth were significantly reduced under drought, with the effect varying across species. Species ranking of drought damage and survival remained consistent with progressing drought. No performance trade-offs emerged between optimal growth and drought resistance of survival (‘growth–stress tolerance’ trade-off hypothesis), or between growth under well-watered and dry conditions (‘growth rates’ trade-off hypothesis). Species local- and large-scale association with moisture (Ellenberg F value and rainfall niche) was not related to their drought resistance. Overall, our results imply that trade-offs and differences of species fundamental drought resistance are not the main drivers of hydrological niche differentiation, species coexistence and their distribution across moisture gradients. The comparative experimental assessment of species whole-plant drought responses we present provides a basis to increase our understanding of current grassland responses to variation of moisture regimes and for projecting consequences of future changes. Electronic supplementary material The online version of this article (10.1007/s00442-020-04625-9) contains supplementary material, which is available to authorized users.

1. Temperate grasslands exhibit strong spatial and temporal variation in water regimes. Thus, grassland plants experience potentially stressful water regimes, which may influence their tissue silicon (Si) and nitrogen (N) concentrations. Plant Si and N concentrations play important ecological roles in temperate grasslands, for example, by influencing plant performance and herbivory, yet comparisons of species' responses to a broad range of water regimes, including drought, waterlogging and flooding, are lacking. 2. We conducted a mesocosm experiment with 10 temperate grassland species of two life-forms (grasses and forbs) exposed to four different soil water regimes (drought, a benign control, waterlogged and flooded conditions), and analysed their Si and N concentrations. 3. Grasses showed lower Si concentrations under drought and flooding compared to the benign control and the highest concentrations emerged under waterlogging. Overall, plant Si responses of grasses were more uniform, while in forbs, responses varied both in direction and magnitude across species. For N concentrations, all species and life-forms showed the highest concentrations under drought compared to the benign control, while half of the species exhibited decreasing concentrations under waterlogging and/or flooding. The water regimes, especially waterlogging and flooding, induced changes in species rankings of plant Si and N concentrations, with stronger shifts in forbs than in grasses. 4. Our results indicate that spatial and temporal variation of water regimes may influence plant Si and N concentrations in temperate grassland species. Plant Si responses to water regimes might be highly species-specific in forbs but more similar in grasses, whereas plant N responses are likely to be relatively uniform across species and life-forms. 5. The strong plasticity in plant Si and N concentrations we observed might have pervasive consequences for ecological processes, such as herbivory.

Understanding how species assembly is influenced by the interplay of climate, local environmental conditions and human-caused disturbances remains a central question in ecology and conservation. Here, we assess how plant species abundance is determined by combinations of functional traits (ecological strategies) and interacting gradients of rainfall, soil conditions (fertility and field capacity) and chronic anthropogenic disturbance in a Caatinga dry tropical forest, Brazil. We tested for trait–environment relationships using multivariate methods (RLQ) accounting for groups of species sharing similar responses to gradients and similar expression of multiple traits (i.e. response groups). Overall, species’ abundances changed predictably in response to rainfall and soil fertility, and were mediated by functional traits, i.e. species with particular trait combinations tended to respond similarly to multifactorial conditions. Briefly, three ecological strategies emerged: species with low wood density and soft (i.e. lower dry matter content), thick leaves converged into a trait syndrome characterizing a drought-avoidance strategy through water storage. They were particularly abundant under extremely low precipitation and relatively high soil field capacity. Under conditions of increasing rainfall and decreasing soil field capacity, species with high wood density were favored, consistent with a drought-tolerance strategy. However, these species fell into two groups relative to leaf-investment: more conservative leaves (low SLA) on relatively fertile soils vs. thinner and softer (i.e. high SLA) leaves on unfertile soils. In seasonally dry tropical forests, low SLA on relatively fertile soils may represent a water conservation strategy. Unexpectedly, no ecological strategy emerged in response to disturbance. The patterns we uncovered help to understand the interplay between precipitation, soil fertility and anthropogenic disturbance in plant species filtering in seasonally dry tropical forests. Moreover, our results underline that impacts of future climate change will depend on how rainfall patterns covary with finer-scale environmental factors such as soil fertility and field capacity.

The plant economics spectrum hypothesizes a correlation among resource-use related traits along one single axis, which determines species’ growth rates and their ecological filtering along resource gradients. This concept has been mostly investigated and shown in perennial species, but has rarely been tested in annual species. Annuals evade unfavorable seasons as seeds and thus may underlie different constraints, with consequences for interspecific trait-trait, trait-growth, and trait-environment relations. To test the hypotheses of the plant economics spectrum in annual species, we measured twelve resource-use related leaf and root traits in 30 winter annuals from Israel under controlled conditions. Traits and their coordinations were related to species’ growth rates (for 19 species) and their distribution along a steep rainfall gradient. Contrary to the hypotheses of the plant economics spectrum, in the investigated annuals traits were correlated along two independent axes, one of structural traits and one of carbon gain traits. Consequently, species’ growth rates were related to carbon gain traits, but independent from structural traits. Species’ distribution along the rainfall gradient was unexpectedly neither associated with species’ scores along the axes of carbon gain or structural traits nor with growth rate. Nevertheless, root traits were related with species’ distribution, indicating that they are relevant for species’ filtering along rainfall gradients in winter annuals. Overall, our results showed that the functional constraints hypothesized by the plant economics spectrum do not apply to winter annuals, leading to unexpected trait-growth and trait-rainfall relations. Our study thus cautions to generalize trait-based concepts and findings between life-history strategies. To predict responses to global change, trait-based concepts should be explicitly tested for different species groups.

Local tree species distributions in tropical forests correlate strongly with soil water availability. However, it is unclear how species distributions are shaped by demographic responses to soil water availability. Specifically, it remains unknown how growth affects species distributions along water availability gradients relative to mortality. 2. We quantified spatial variation in dry season soil water potential (SWP) in the moist tropical forest on Barro Colorado Island, Panama, and used a hierarchical Bayesian approach to evaluate relationships between demographic responses of naturally regenerating seedlings to SWP (RGRs and first‐year mortality) and species distributions along the SWP gradient for 62 species. We also tested whether species that were more abundant at the wet or dry end of the gradient performed better (a) at their “home end” of the gradient (“best at home” hypothesis) and (b) “at home” compared to co‐occurring species (“home advantage” hypothesis). 3. Four and five species responded significantly to SWP in terms of growth or mortality respectively. Growth (but not mortality) responses were positively related to species distributions along the SWP gradient; species with a more positive (negative) growth response to SWP were more abundant at higher (lower) SWP, that is, at wetter (drier) sites. In addition, wet distributed species grew faster on the wet end of the SWP gradient than on the dry end (“best at home”) and grew faster on the wet end than dry distributed species (“home advantage”). Mortality rates declined with seedling size for all species. Thus, seedling growth responses to SWP indirectly shaped local species distributions by influencing seedling size and thereby mortality risk. 4. Synthesis. By demonstrating how growth responses to spatial variation in soil water availability affect species distributions, we identified a demographic process underlying niche differentiation on hydrological gradients in tropical forests. Recognizing the role of these growth responses in shaping species distributions should improve the understanding of tropical forest composition and diversity along rainfall gradients and with climate change.

Silicon (Si) has a variety of functions in plants, including alleviation of drought and light stress, defense against herbivores and pathogens, and improving plant nutrition. However, for tropical forests our knowledge about the role of silicon and its variation in soils and plants remains limited. To advance our insights into the potential role of Si in tropical forest ecology, we combined observational and experimental approaches to assess (i) variation in soluble and amorphous Si concentrations in tropical forest soils at the local and regional scale, and their relation to soil weathering stage, soil chemistry, and rainfall, (ii) variation of foliar Si concentrations across more than 30 co-occurring woody species, and (iii) intra-specific variation of foliar Si across sites and foliar habits (sun and shade). We found considerable (27-fold) variation in soluble Si (extracted in 0.01 M CaCl2) across soils, which reflected soil weathering stage and chemistry, but not rainfall. Foliar Si also varied markedly across species, both in naturally occurring trees and in a common garden experiment, with 39% of the most abundant species being classified as Si accumulators. Within species, foliar Si varied among sites and foliar habits, but relationships were inconsistent across species. The marked variability of Si concentrations in soils and leaves indicates that Si is likely to play an important yet underappreciated role for a variety of ecological processes in tropical forests.

Seedlings in moist tropical forests must cope with deep shade and seasonal drought. However, the interspecific relationship between seedling performance in shade and drought remains unsettled. We quantified spatiotemporal variation in shade and drought in the seasonal moist tropical forest on Barro Colorado Island (BCI), Panama, and estimated responses of naturally regenerating seedlings as the slope of the relationship between performance and shade or drought intensity. Our performance metrics were relative height growth and first-year survival. We investigated the relationship between shade and drought responses for up to 63 species. There was an interspecific trade-off in species responses to shade versus species responses to dry season intensity; species that performed worse in the shade did not suffer during severe dry seasons and vice versa. This trade-off emerged in part from the absence of species that performed particularly well or poorly in both drought and shade. If drought stress in tropical forests increases with climate change and as solar radiation is higher during droughts, the trade-off may reinforce a shift towards species that resist drought but perform poorly in the shade by releasing them from deep shade.

In tropical forests, insect herbivores exert significant pressure on plant populations. Adaptation to such pressure is hypothesized to be a driver of high tropical diversity, but direct evidence for local adaptation to herbivory in tropical forests is sparse. At the same time, herbivore pressure has been hypothesized to increase with rainfall in the tropics, which could lead to differences among sites in the degree of local adaptation. To assess the presence of local adaptation and its interaction with rainfall, we compared herbivore damage on seedlings of local vs. nonlocal populations at sites differing in moisture availability in a reciprocal transplant experiment spanning a rainfall gradient in Panama. For 13 native tree species, seeds collected from multiple populations along the rainfall gradient were germinated in a shadehouse and then transplanted to experimental sites within the species range. We tracked the proportion of seedlings attacked over 1.5 yr and quantified the percentage of leaf area damaged at the end of the study. Seedlings originating from local populations were less likely to be attacked and experienced lower amounts of herbivore damage than those from nonlocal populations, but only on the wetter end of the rainfall gradient. However, overall herbivore damage was higher at the drier site compared to wetter sites, contrary to expectation. Taken together, these findings support the idea that herbivory can result in local adaptation within tropical tree species; however, the likelihood of local adaptation varies among sites because of environmentally driven differences in investment in defense or herbivore specialization or both.

Hydraulic traits are important for woody plant functioning and distribution. Associations among hydraulic traits, other leaf and stem traits, and species’ performance are relatively well understood for trees, but remain poorly studied for lianas. We evaluated the coordination among hydraulic efficiency (i.e., maximum hydraulic conductivity), hydraulic safety (i.e., cavitation resistance), a suite of eight morphological and physiological traits, and species’ abundances for saplings of 24 liana species and 27 tree species in wet tropical forests in Panama. Trees showed a strong trade‐off between hydraulic efficiency and hydraulic safety, whereas efficiency and safety were decoupled in lianas. Hydraulic efficiency was strongly and similarly correlated with acquisitive traits for lianas and trees (e.g., positively with gas exchange rates and negatively with wood density). Hydraulic safety, however, showed no correlations with other traits in lianas, but with several in trees (e.g., positively with leaf dry matter content and wood density and negatively with gas exchange rates), indicating that in lianas hydraulic efficiency is an anchor trait because it is correlated with many other traits, while in trees both efficiency and safety are anchor traits. Traits related to shade tolerance (e.g., low specific leaf area and high wood density) were associated with high local tree sapling abundance, but not with liana abundance. Our results suggest that different, yet unknown mechanisms determine hydraulic safety and local‐scale abundance for lianas compared to trees. For trees, the trade‐off between efficiency and safety will provide less possibilities for ecological strategies. For lianas, however, the uncoupling of efficiency and safety could allow them to have high hydraulic efficiency, and hence high growth rates, without compromising resistance to cavitation under drought, thus allowing them to thrive and outperform trees under drier conditions.

One explanation for the extraordinarily high tree diversity of tropical lowland forests is that it is maintained by specialized natural enemies such as insect herbivores, which cause distance- and density-dependent mortality. Insect herbivory could also explain the positive correlation between tree species richness and rainfall if herbivory increases with rainfall, is higher on locally abundant versus rare species, and is not limited by predation pressure at wet sites. To test these predictions, insect herbivory and predation pressure on insect herbivores were quantified across a neotropical rainfall and tree species richness gradient, and herbivory was investigated in relation to local tree abundances. Insect herbivory on leaves (folivory) decreased strongly and significantly with rainfall, while predation pressure was significantly higher at the wetter site. Herbivores were more likely to attack abundant tree species, but herbivore damage levels were not related to tree species abundance. Insect folivores might contribute to local tree species coexistence in our system, but seem unlikely to drive the positive correlation between tree species richness and rainfall. The unexpected and contrasting patterns of herbivory and predation we observed support the need for a multi-trophic perspective to understand fully the processes contributing to diversity and ecosystem functioning.

1. Forest dieback caused by drought-induced tree mortality has been observed world-wide. Forecasting which trees in which locations are vulnerable to drought-induced mortality is important to predict the consequences of drought on forest structure, biodiversity and ecosystem function. 2. In this paper, our central aim was to compile a synthesis of tree traits and associated abiotic variables that can be used to predict drought-induced mortality. We reviewed the literature that specifically links drought mortality to functional traits and site conditions (i.e. edaphic variables and biotic conditions), targeting studies that show clear use of tree traits in drought analysis. We separated the review into five climatic zones to determine global vs. regionally restricted relationships between traits and mortality. 3. Our synthesis identifies a number of traits that have clear relationships with drought-induced mortality (e.g. wood density at the species level and tree size and growth at the individual level). However, the lack of direct relationships between most traits and drought-induced mortality highlights areas where future research should focus to broaden our understanding. 5. Synthesis and applications. Our synthesis highlights established relationships between traits and drought-induced mortality, presents knowledge gaps for future research focus and suggests monitoring and research avenues for improving our understanding of drought-induced mortality. It is intended to assist ecologists and natural resource managers choose appropriate and measurable parameters for predicting local and regional scale tree mortality risk in different climatic zones within constraints of time and funding availability.

Phosphorus is widely considered to constrain primary productivity in tropical rain forests, yet the chemistry of soil organic phosphorus in such ecosystems remains poorly understood. We assessed the composition of soil organic phosphorus in 19 contrasting soils under lowland tropical forest in the Republic of Panama using NaOH–EDTA extraction and solution 31P nuclear magnetic resonance spectroscopy. The soils spanned a strong rainfall gradient (1730–3404 mm y−1) and contained a wide range of chemical properties (pH 3.3–7.0; total carbon 2.8–10.4%; total phosphorus 74–1650 mg P kg−1). Soil organic phosphorus concentrations ranged between 22 and 494 mg P kg−1 and were correlated positively with total soil phosphorus, pH, and total carbon, but not with annual rainfall. Organic phosphorus constituted 26 ± 1% (mean ± STD error, n = 19) of the total phosphorus, suggesting that this represents a broad emergent property of tropical forest soils. Organic phosphorus occurred mainly as phosphate monoesters (68–96% of total organic phosphorus) with smaller concentrations of phosphate diesters in the form of DNA (4–32% of total organic phosphorus). Phosphonates, which contain a direct carbon–phosphorus bond, were detected in only two soils (3% of the organic phosphorus), while pyrophosphate, an inorganic polyphosphate with a chain length of two, was detected in all soils at concentrations up to 13 mg P kg−1 (3–13% of extracted inorganic phosphorus). Phosphate monoesters were a greater proportion of the total organic phosphorus in neutral soils with high concentrations of phosphorus and organic matter, whereas the proportion of phosphate diesters was greater in very acidic soils low in phosphorus and organic matter. Most soils did not contain detectable concentrations of either myo- or scyllo-inositol hexakisphosphate, which is in marked contrast to many temperate mineral soils that contain abundant inositol phosphates. We conclude that soil properties exert a strong control on the amounts and forms of soil organic phosphorus in tropical rain forests, but that the proportion of the total phosphorus in organic forms is relatively insensitive to variation in climate and soil properties. Further work is now required to assess the contribution of soil organic phosphorus to the nutrition and diversity of plants in these species-rich ecosystems.

Summary 1. Habitat specialization models predict that adaptations to environmental conditions explain species distributions. In tropical rainforests, the ability of the seedlings to survive during drought has been shown to be a key determinant of species distributions. We hypothesize that differences among species in their tolerance to low tissue water status is the mechanism underlying differences in performance during drought. 2. To test this hypothesis we quantified tolerance to low leaf water status for over 20 species from central Panama in screenhouse experiments using two different experimental approaches. Results from both approaches were highly correlated with each other. 3. We found that tolerance to low leaf water status correlated with species drought performance in the field and with their distribution across a gradient of dry season length, with the more desiccationtolerant species having higher survival in dry relative to irrigated conditions, and occurring in drier areas. These results support the hypothesis that, in tropical forests, tolerance to low tissue water status governs seedling performance during drought, as well as being a determinant of species distribution patterns. 4. Lower tolerance to low leaf water status was correlated with greater stem hydraulic conductance. In addition, all species tested, including both desiccation-sensitive and desiccation-resistant species, showed similar losses of xylem conductance, about 80%, when near death. These results suggest that a principal mechanism by which desiccation leads to plant mortality is the loss of xylem conductivity.

Rainfall and soil moisture availability vary greatly both spatially and temporally. They are prime factors influencing species distribution patterns, diversity and habitat associations in a range of biomes, and limit primary productivity in many natural ecosystems, as well as in forestry and agricultural systems (Hawkins et al. 2003, Kozlowski & Pallardy 1997, Lieth 1975). Projections of drying trends, and increased frequency and intensity of drought events with climate and land-use changes (Hulme & Viner 1998, IPCC 2001) have fuelled an increased interest in the role of drought in determining the structure and function of natural and managed forest systems (Allen & Breshears 1998, Miles et al. 2004). Such projections accentuate the need to assess, understand and predict plant reactions to drought, as well as soil moisture variation at different scales.

Drought strongly influences plant phenology, growth and mortality in tropical forests, thereby shaping plant performance, population dynamics and community structure (Bunker & Carson 2005, Condit et al. 1995). Microbial symbionts of plants profoundly influence host water relations (Lösch & Gansert 2002), but are rarely considered in studies of tropical plant physiology. In particular, plant–fungus associations, which are ubiquitous in plant communities and especially common in tropical forests, play important and varied roles in plant water status. Fungal pathogens associated with roots, vascular tissue and foliage may interfere with water uptake and transport, increase rates of foliar transpiration, and induce xylem embolism and tissue death (Agrios 1997). In contrast, rhizosphere mutualists such as ecto- and arbuscular mycorrhizal fungi may benefit hosts by increasing surface area for water uptake, enhancing stomatal regulation of water loss, and increasing root hydraulic conductivity (Auge 2001, Lösch & Gansert 2002).

We used regression models and information-theoretic model selection to assess the relative importance of environment, local dispersal and historical contingency as controls of the distributions of 26 common plant species in tropical forest on Barro Colorado Island (BCI), Panama. We censused eighty-eight 0.09-ha plots scattered across the landscape. Environmental control, local dispersal and historical contingency were represented by environmental variables (soil moisture, slope, soil type, distance to shore, old-forest presence), a spatial autoregressive parameter (ρ), and four spatial trend variables, respectively. We built regression models, representing all combinations of the three hypotheses, for each species. The probability that the best model included the environmental variables, spatial trend variables and ρ averaged 33%, 64% and 50% across the study species, respectively. The environmental variables, spatial trend variables, ρ, and a simple intercept model received the strongest support for 4, 15, 5 and 2 species, respectively. Comparing the model results to information on species traits showed that species with strong spatial trends produced few and heavy diaspores, while species with strong soil moisture relationships were particularly drought-sensitive. In conclusion, history and local dispersal appeared to be the dominant controls of the distributions of common plant species on BCI.

We investigated how species identity and variation in salinity and nutrient availability influence the hydraulic conductivity of mangroves. Using a fertilization study of two species in Florida, we found that stem hydraulic conductivity expressed on a leaf area basis (K-leaf) was significantly different among species of differing salinity tolerance, but was not significantly altered by enrichment with limiting nutrients. Reviewing data from two additional sites (Panama and Belize), we found an overall pattern of declining leaf-specific hydraulic conductivity (K-leaf) with increasing salinity. Over three sites, a general pattern emerges, indicating that native stem hydraulic conductivity (K-h) and K-leaf are less sensitive to nitrogen (N) fertilization when N limits growth, but more sensitive to phosphorus (P) fertilization when P limits growth. Processes leading to growth enhancement with N fertilization are probably associated with changes in allocation to leaf area and photosynthetic processes, whereas water uptake and transport processes could be more limiting when P limits growth. These findings suggest that whereas salinity and species identity place broad bounds on hydraulic conductivity, the effects of nutrient availability modulate hydraulic conductivity and growth in complex ways.

Spatial gradients in mangrove tree height in barrier islands of Belize are associated with nutrient deficiency and sustained flooding in the absence of a salinity gradient. While nutrient deficiency is likely to affect many parameters, here we show that addition of phosphorus (P) to dwarf mangroves stimulated increases in diameters of xylem vessels, area of conductive xylem tissue and leaf area index (LAI) of the canopy. These changes in structure were consistent with related changes in function, as addition of P also increased hydraulic conductivity (K-s), stomatal conductance and photosynthetic assimilation rates to the same levels measured in taller trees fringing the seaward margin of the mangrove. Increased xylem vessel size and corresponding enhancements in stern hydraulic conductivity in P fertilized dwarf trees came at the cost of enhanced midday loss of hydraulic conductivity and was associated with decreased assimilation rates in the afternoon. Analysis of trait plasticity identifies hydraulic properties of trees as more plastic than those of leaf structural and physiological characteristics, implying that hydraulic properties are key in controlling growth in mangroves. Alleviation of P deficiency, which released trees from hydraulic limitations, reduced the structural and functional distinctions between dwarf and taller fringing tree forms of Rhizophora mangle.

Plant productivity, distribution and diversity in tropical rain forests correlate with water availability. Water availability is determined by rainfall and also by the available water capacity of the soil. However, while rainfall is recognized as important, linkages between plant distribution and differences among soils in available water capacity have not been demonstrated. One reason for this may be that measurements of soil moisture, such as gravimetric water content, may be overly simplistic. To investigate this, we compared two sites in Panama, Allee and Rio Paja, which have similar rainfall but different plant communities. Soil water release curves were obtained from about −0.1 MPa to −9 MPa, permitting us to calculate available water capacity. The Rio Paja site had 17% greater available water capacity (between −0.1 MPa to −3 MPa), whereas the gravimetric water content at Rio Paja was lower by 16% in rainy season and by 41% at the end of the dry season. Hence soil gravimetric water content and soil available water capacity did not correspond. The results suggest that available water capacity may better predict plant distributions. Hence, whenever possible, available water capacity should be determined in addition to gravimetric water content.

The amount and seasonality of rainfall varies strongly in the tropics, and plant species abundance, distribution and diversity are correlated with rainfall. Drought periods leading to plant stress occur not only in dry forests, but also in moist and even wet forests. We quantified experimentally the effect of drought on survival of first year seedlings of 28 co-occurring tropical woody plant species in the understory of a tropical moist forest. The seedlings were transplanted to plots and subjected to a drought and an irrigation treatment for 22 weeks during the dry season. Drought effects on mortality and wilting behavior varied greatly among species, so that relative survival in the dry treatment ranged from 0% to about 100% of that in the irrigated treatment. Drought stress was the main factor in mortality, causing about 90% (median) of the total mortality observed in the dry treatment. In almost half of the species, the difference in survival between treatments was not significant even after 22 weeks, implying that many of the species are well adapted to drought in this forest. Relative drought survival was significantly higher in species associated with dry habitats than in those associated with wet habitats, and in species with higher abundance on the dry side of the Isthmus of Panama, than in those more abundant on the wet side. These data show that differential species survival in response to drought, combined with variation in soil moisture availability, may be important for species distribution at the local and regional scale in many tropical forests.

The ecological mechanisms hypothesized to structure species-rich communities range from strict local determinism to neutral ecological drift. We assessed the degree of ecological determinism in tropical plant community structure by analyses of published demographic data; a broad range of spatial, historical, and environmental variables; and the distributions of 33 herbaceous species (plot size = 0.02 ha) and 61 woody species (plot size = 0.09 ha) among 350 plots in a 16-km2 forest landscape (Barro Colorado Island, Panama). We found a strong degree of cross-landscape dominance by a subset of species whose identities were predictable from sapling survivorship rates under shade. Using canonical ordination we found that spatial and environmental–historical factors were of comparable importance for controlling within-landscape variability in species composition. Past land use had a strong impact on species composition despite ceasing 100–200 years ago. Furthermore, edaphic–hydrological factors, treefall gaps, and an edge effect all had unique impacts on species composition. Hence, ecological determinism was evident in terms of both cross-landscape dominance and within-landscape variability in species composition. However, at the latter scale, the large portion of the explained variance in species composition among plots uniquely attributed to spatial location pointed to an equally important role for neutral processes.

Quantifying plant drought resistance is important for understanding plant species' association to microhabitats with different soil moisture availability and their distribution along rainfall gradients, as well as for understanding the role of underlying morphological and physiological mechanisms. The effect of dry season drought on survival and leaf-area change of first year seedlings of 28 species of co-occurring woody tropical plants was experimentally quantified in the understory of a tropical moist forest. The seedlings were subjected to a drought or an irrigation treatment in the forest for 22 weeks during the dry season. Drought decreased survival and growth (assessed as leaf-area change) in almost all of the species. Both survival and leaf-area change in the dry treatment ranged fairly evenly from 0% to about 100% of that in the irrigated treatment. In 43% of the species the difference between treatments in survival was not significant even after 22 weeks. In contrast, only three species showed no significant effect of drought on leaf-area change. The effects of drought on species' survival and growth were not correlated with each other, reflecting different strategies in response to drought. Seedling size at the onset of the dry season had no significant effect on species' drought response. Our study is the first to comparatively assess seedling drought resistance in the habitat for a large number of tropical species, and underlines the importance of drought for plant population dynamics in tropical forests.

Studies of the desiccation tolerance of the seedlings of five tropical trees were made on potted plants growing in a greenhouse. Pots were watered to field capacity and then dehydrated for 3 to 9 weeks to reach various visual wilting stages, from slightly wilted to dead. Saturated root hydraulic conductance was measured with a high-pressure flowmeter, and whole-stem hydraulic conductance was measured by a vacuum chamber method. Leaf punches (5.6-mm diameter) were harvested for measurement of leaf water potential by a thermocouple psychrometer method and for measurement of fresh and dry weight. In a parallel study, the same five species were studied in a field experiment in the understory of a tropical forest, where these species frequently germinate. Control seedlings were maintained in irrigated plots during a dry season, and experimental plants were grown in similar plots with rain exclusion shelters. Every 2 to 4 weeks, the seedlings were scored for wilt state and survivorship. After a 22-week drought, the dry plots were irrigated for several weeks to verify visual symptoms of death. The field trials were used to rank drought performance of species, and the greenhouse desiccation studies were used to determine the conditions of moribund plants. Our conclusion is that the desiccation tolerance of moribund plants correlated with field assessment of drought-performance for the five species (r 2 > 0.94).

In tropical forests, severe droughts caused by El Niño events may strongly influence the water relations of tree seedlings and thereby increase their mortality. Data on known-aged seedlings of three common shade-tolerant canopy tree species (Trichilia tuberculata, Tetragastris panamensis and Quararibea asterolepis) in a Panamanian moist forest are presented. Seedling survival during a severe El Niño dry season (1997-98) was compared with prior long-term survival data, and levels of drought stress were assessed by measuring plant water potentials and gas exchange characteristics. Contrary to prediction, dry-season seedling survival was not dramatically reduced in any species compared with that expected in ‘normal’ years. In Trichilia and Quararibea, pre-dawn water potentials averaged −2 MPa and midday water potentials about −3 MPa. Stomatal conductances were very low, averaging 26 mmol m-2 s-1 for Tetragastris and 11–13 mmol m-2 s-1 for Trichilia and Quararibea. Photosynthetic rates also were very low but consistently positive, averaging 0.8–1.1 μmol m-2 s-1. The findings suggest that, once established, seedlings of common tree species in this semi-deciduous forest may be tolerant of drought events.

The suitability of several methods for estimating light conditions in the understorey of tropical forests, and of different sampling schedules was evaluated. Light conditions at 16 understorey sites in a Panamanian lowland forest were continuously measured for 9 mo with quantum sensors and photodiodes. Light conditions at the sites were also assessed indirectly with hemispherical fisheye photographs, plant canopy analysis, 38-mm photographs, 24-mm photographs and a spherical densiometer. Estimates from all indirect methods, except the spherical densiometer, were highly correlated with the direct measurements. Short-term direct light measurements for a day or a week also correlated with long-term light conditions. The indirect measures differed by up to c. 70% from the direct measures relative to single site measurements. Hence, the indirect methods are inadequate where single site light conditions have to be assessed accurately. However, because light conditions encountered in the understorey varied up to 13-fold, the indirect methods were found to be well suited to rank understorey light conditions among a large number of sites. The results from frequent and infrequent sampling schedules differed only slightly, suggesting that taking indirect measures at the beginning and the end of a study offers a reasonable compromise between accuracy and sampling effort.

Kalanchoe pinnata (Lam.) Pers. (Crassulaceae), a succulent-leaved crassulacean-acid-metabolism plant, was grown in open-top chambers at ambient and elevated (two times ambient) CO2 concentrations under natural conditions at the Smithsonian Tropical Research Institute, Republic of Panama. Nocturnal increase in titratable acidity and nocturnal carbon gain were linearly related, increased with leaf age, and were unaffected by CO2 treatments. However, under elevated CO2, dry matter accumulation increased by 42–51%. Thus, the increased growth at elevated CO2 was attributable entirely to increased net CO2 uptake during daytime in the light. Malic acid was the major organic acid accumulated overnight. Nocturnal malate accumulation exceeded nocturnal citrate accumulation by six-to eightfold at both CO2 concentrations. Basal (predawn) starch levels were higher in leaves of plants grown at elevated CO2 but diurnal fluctuations of starch were of similar magnitude under both ambient and elevated CO2. In both treatments, nocturnal starch degradation accounted for between 78 and 89% of the nocturnal accumulation of malate and citrate. Glucose, fructose, and sucrose were not found to exhibit marked day-night fluctuations.

Tropical forest vegetation is shaped by climate and by soil, but understanding how the distributions of individual tree species respond to specific resources has been hindered by high diversity and consequent rarity. To study species over an entire community, we surveyed trees and measured soil chemistry across climatic and geological gradients in central Panama and then used a unique hierarchical model of species occurrence as a function of rainfall and soil chemistry to circumvent analytical difficulties posed by rare species. The results are a quantitative assessment of the responses of 550 tree species to eight environmental factors, providing a measure of the importance of each factor across the entire tree community. Dry-season intensity and soil phosphorus were the strongest predictors, each affecting the distribution of more than half of the species. Although we anticipated clear-cut responses to dry-season intensity, the finding that many species have pronounced associations with either high or low phosphorus reveals a previously unquantified role for this nutrient in limiting tropical tree distributions. The results provide the data necessary for understanding distributional limits of tree species and predicting future changes in forest composition.

Throughout the evolutionary history of plants, drought, shade, and scarcity of nutrients have structured ecosystems and communities globally. Humans have begun to drastically alter the prevalence of these environmental factors with untold consequences for plant communities and ecosystems worldwide. Given limitations in using organ-level traits to predict ecological performance of species, recent advances using tolerances of low resource availability as plant functional traits are revealing the often hidden roles these factors have in structuring communities and are becoming central to classifying plants ecologically. For example, measuring the physiological drought tolerance of plants has increased the predictability of differences among species in their ability to survive drought as well as the distribution of species within and among ecosystems. Quantifying the shade tolerance of species has improved our understanding of local and regional species diversity and how species have sorted within and among regions. As the stresses on ecosystems continue to shift, coordinated studies of whole-plant growth centered on tolerance of low resource availability will be central in predicting future ecosystem functioning and biodiversity. This will require efforts that quantify tolerances for large numbers of species and develop bioinformatic and other techniques for comparing large number of species.

Summary Short-term responses of net CO 2 assimilation rate (A), in the light and dark, to ambient CO 2 partial pressure (between about 30 and 1000 µbar) were studied in leaves of the crassulacean acid metabolism (CAM) plant, Kalanchoe pinnata . The results show that it is possible to extrapolate from instantaneous measurements of net CO 2 exchange to diurnal and nocturnal CO 2 balances at different CO 2 partial pressures. Instantaneous CO 2 response curves were obtained by altering CO 2 levels at 10-min intervals during the middle of the 12-h dark period (phase I of CAM) and during the last third of the 12-h light period (phase IV of CAM). CO 2 partial pressures were also altered at longer, 12- to 24-h intervals and maximum rates of net CO 2 uptake (A max ) during light and dark periods were analysed in response to intercellular CO 2 partial pressures (p i ) occurring at the time of A max· A max-Pi relationships were identical to A-p i -curves from rapidly performed determinations during phases I and IV. Corresponding to previous findings with non-CAM species, A max and integrated net carbon gain during light and dark periods, respectively, showed a linear relationship. Nocturnal CO 2 uptake in normal air was barely affected when lightperiod carbon gain was manipulated by alterations in CO 2 partial pressure. Carbon gain during light periods, measured in normal air, was also independent of CO 2 -related changes in nocturnal carbon gain. Only after 12 h of darkness at the lowest CO 2 concentration used (about 30 µbar), was carbon gain increased under lighted conditions.

A trade-off between growth and mortality rates characterizes tree species in closed canopy forests. This trade-off is maintained by inherent differences among species and spatial variation in light availability caused by canopy-opening disturbances. We evaluated conditions under which the trade-off is expressed and relationships with four key functional traits for 103 tree species from Barro Colorado Island, Panama. The trade-off is strongest for saplings for growth rates of the fastest growing individuals and mortality rates of the slowest growing individuals (r 2 ¼ 0.69), intermediate for saplings for average growth rates and overall mortality rates (r 2 ¼ 0.46), and much weaker for large trees (r 2 � 0.10). This parallels likely levels of spatial variation in light availability, which is greatest for fast- vs. slow-growing saplings and least for large trees with foliage in the forest canopy. Inherent attributes of species contributing to the trade-off include abilities to disperse, acquire resources, grow rapidly, and tolerate shade and other stresses. There is growing interest in the possibility that functional traits might provide insight into such ecological differences and a growing consensus that seed mass (SM), leaf mass per area (LMA), wood density (WD), and maximum height (Hmax) are key traits among forest trees. Seed mass, LMA, WD, and Hmax are predicted to be small for light-demanding species with rapid growth and mortality and large for shade-tolerant species with slow growth and mortality. Six of these trait-demographic rate predictions were realized for saplings; however, with the exception of WD, the relationships were weak (r 2 , 0.1 for three and r 2 , 0.2 for five of the six remaining relationships). The four traits together explained 43-44% of interspecific variation in species positions on the growth-mortality trade-off; however, WD alone accounted for .80% of the explained variation and, after WD was included, LMA and Hmax made insignificant contributions. Virtually the full range of values of SM, LMA, and Hmax occurred at all positions on the growth-mortality trade-off. Although WD provides a promising start, a successful trait- based ecology of tropical forest trees will require consideration of additional traits.

Associations with topographic units or soil types that vary in water availability are widespread in plant communities and are one of the main structuring aspects for local species distribution patterns, yet the underlying mechanisms are poorly understood. We hypothesized that differential seedling performance across habitats, particularly during the dry season, leads to habitat associations in seasonal tropical forests. We expected this pattern to be most pronounced in particularly dry years, such as those associated with El Nino Southern Oscillation (ENSO) events. We assessed performance of 36 native tree and shrub species in a moist forest in central Panama across the dry and wet seasons in two topographic habitat types during a year in which dry-season precipitation was reduced due to an ENSO event. At the community level, we found lower growth and higher mortality in the dry season relative to the wet season and higher mortality in the drier plateau habitat relative to the wetter slope habitat. There was large variation in species' responses to season and habitat. Species' mortality and growth rates were significantly correlated with experimentally assessed drought sensitivity, but only during the dry season in the plateau habitat. Slope specialists had significantly higher survival, but not growth, in the slope vs. plateau habitat during the dry season. In contrast, plateau specialists showed no performance differences between habitats. The data suggest that associations with plateau habitats result from a numerical advantage of drought-tolerant species in dry habitats in which seedlings of drought-sensitive species are unable to persist. Overall, our results support the idea that seasonal and spatial variation in water availability, particularly in dry years, drive seedling dynamics, which in turn shape local species distributions. Predicted shifts in rainfall patterns caused by global and regional climate change are therefore expected to alter the dynamics, composition, and diversity of seasonal tropical forests.

In moist tropical forests resprouting may be an important component of life history, contributing to asexual reproduction through the clonal spread of individuals derived from shoot fragments. However, in contrast to other ecosystems where resprouting is common, the ecological correlates of resprouting capacity in tropical forests remain largely unexplored. In this study we characterized shade tolerance, resprouting capacity and sexual reproductive success of eight co-occurring Piper species from lowland forests of Panama. In field experiments we found that shade-tolerant Piper species had a higher capacity to regenerate from excised or pinned stem fragments than light-demanding species in both gap and understory light conditions. In contrast, shade-tolerant species had lower recruitment probabilities from seeds, as a consequence of lower initial seed viability, and lower seedling emergence rates. All Piper species needed gap conditions for successful seedling establishment. Of 8,000 seeds sown in the understory only 0.2% emerged. In gaps, seed germination of light-demanding species was between 10 and 50%, whereas for shade-tolerant species it was 0.5-9.8%. We propose that the capacity to reproduce asexually from resprouts could be adaptive for shade-tolerant species that are constantly exposed to damage from falling litter in the understory. Resprouting may allow Piper populations to persist and spread despite the high rate of pre-dispersal seed predation and low seed emergence rates. Across Piper species, we detected a trade-off between resprouting capacity and the annual viable seed production per plant but not with annual seed mass produced per plant. This suggests that species differences in sexual reproductive success may not necessarily result from differential resource allocation. Instead we suggest that low sexual reproductive success in the understory may in part reflect reduced genetic diversity in populations undergoing clonal growth, resulting in self-fertilization and in-breeding depression.

Mangrove ecosystems can be either nitrogen (N) or phosphorus (P) limited and are therefore vulnerable to nutrient pollution. Nutrient enrichment with either N or P may have differing effects on ecosystems because of underlying differences in plant physiological responses to these nutrients in either N- or P-limited settings. Using a common mangrove species, Avicennia germinans, in sites where growth was either N or P limited, we investigated differing physiological responses to N and P limitation and fertilization. We tested the hypothesis that water uptake and transport, and hydraulic architecture, were the main processes limiting productivity at the P-limited site, but that this was not the case at the N-limited site. We found that plants at the P-deficient site had lower leaf water potential, stomatal conductance and photosynthetic carbon-assimilation rates, and less conductive xylem, than those at the N-limited site. These differences were greatly reduced with P fertilization at the P-limited site. By contrast, fertilization with N at the N-limited site had little effect on either photosynthetic or hydraulic traits. We conclude that growth in N- and P-limited sites differentially affect the hydraulic pathways of mangroves. Plants experiencing P limitation appear to be water deficient and undergo more pronounced changes in structure and function with relief of nutrient deficiency than those in N-limited ecosystems.

Variation in plant species performance in re- sponse to water availability offers a potential axis for temporal and spatial habitat partitioning and may therefore affect community composition in tropical for- ests. We hypothesized that short dry spells during the wet season are a significant source of mortality for the newly emerging seedlings of pioneer species that recruit in treefall gaps in tropical forests. An analysis of a 49- year rainfall record for three forests across a rainfall gradient in central Panama confirmed that dry spells of ‡10 days during the wet season occur on average once a year in a deciduous forest, and once every other year in a semi-deciduous moist and an evergreen wet forest. The effect of wet season dry spells on the recruitment of pioneers was investigated by comparing seedling sur- vival in rain-protected dry plots and irrigated control plots in four large artificially created treefall gaps in a semi-deciduous tropical forest. In rain-protected plots surface soil layers dried rapidly, leading to a strong gradient in water potential within the upper 10 cm of soil. Seedling survival for six pioneer species was sig- nificantly lower in rain-protected than in irrigated con- trol plots after only 4 days. The strength of the irrigation effect differed among species, and first became apparent 3-10 days after treatments started. Root allocation patterns were significantly, or marginally significantly, different between species and between two groups of larger and smaller seeded species. However, they were not correlated with seedling drought sensitivity, sug- gesting allocation is not a key trait for drought sensi- tivity in pioneer seedlings. Our data provide strong evidence that short dry spells in the wet season differ- entially affect seedling survivorship of pioneer species, and may therefore have important implications to seedling demography and community dynamics.

An analysis of the physiological vulnerability of different trees to drought shows that forests around the globe are at equally high risk of succumbing to increases in drought conditions. See Letter p.752 Forest dieback resulting from extreme drought events has the potential to cause widespread loss of biodiversity, with a major impact on the global carbon balance. Plants undergoing drought stress experience reduced xylem pressure, and each species can tolerate a different degree of reduction before xylem damage and, eventually, hydraulic failure occur. This study looks at the safety margin between minimum experienced xylem pressure and the damage threshold for 226 forest species from 81 sites worldwide and shows that most species across both dry and wet biomes have small safety margins against injurious levels of drought stress. These plants are potentially vulnerable to the combination of rising temperatures and declining rainfall that is predicted to cause droughts of increasing intensity and duration in the near future.

Studies of the desiccation tolerance of 15-month-old Licania platypus (Hemsl.) Fritsch seedlings were performed on potted plants. Pots were watered to field capacity and then dehydrated for 23-46 d to reach various visible wilting stages from slightly-wilted to dead. Root hydraulic conductance, k(r), was measured with a high-pressure flow meter and whole-stem hydraulic conductance, k(ws), was measured by a vacuum chamber method. Leaf punches were harvested for measurement of leaf water potential by a thermocouple psychrometer and for measurement of fresh- and dry-weight. L. platypus was surprisingly desiccation-tolerant, suggesting that most species of central Panama may be well adapted to the seasonality of rainfall in the region. The slightly-wilted stage corresponded to leaf water potentials and relative water contents of -2.7 MPa and 0.85, respectively, but plants did not die until these values fell to -7.5 MPa and 0.14, respectively. As desiccation proceeded k(r) and k(ws) declined relative to irrigated controls, but k(ws) was more sensitive to desiccation than k(r). Values of k(ws) declined by 70-85% in slightly-wilted to dead plants, respectively. By comparison, k(r) showed no significant change in slightly-wilted plants and fell by about 50% in plants having severely-wilted to dead shoots.

International audience; Whole plant hydraulic conductance was measured for two co-occuring neotropical rainforest understory shrub species of the genus Piper, which differ in their habitat preference with respect to soil moisture: P. trigonum is restricted to wet habitats, whereas P. cordulatum occurs in drier sites. Plants were grown under four treatments (two light conditions crossed with two watering regimes), representing the extremes of conditions in their natural habitat. The data showed significantly higher total hydraulic conductance and leaf specific conducance in the drought-avoiding species, P. trigonum, than in the drought-tolerant species P. cordulatum. The measured parameters also differed between growth treatments. The species differences in hydraulic conductance could be interpreted as adaptive in the respective species' habitat, and might thus be important in determining habitat preference and distribution of the two species.; Conductance hydraulique de deux arbustes néotropicaux du sous étage ayant des habitats préférentiels. La conductance hydraulique de la plante entière a été mesurée pour deux espèces arbustives du genre Piper du sous étage d'une forêt néotropicale. Ces espèces différent par leur adaptation à l'humidité du sol : P. trigonum est localisé dans les habitats humides, alors que P. cordulatum se rencontre dans les zones sèches. Les plants ont été soumis à quatre traitements (deux conditions d'éclairement combinées avec deux régimes hydriques), représentant les conditions extrêmes de leurs habitats naturels. Les résultats montrent une conductance hydraulique et une conductance spécifique des feuilles significativement plus élevées pour l'espèce évitant la sécheresse, P. trigonum, par rapport à l'espèce tolérant la sécheresse, P. cordulatum. Les paramètres de croissance mesurés diffèrent aussi selon les traitements. Les différences de conductance hydraulique entre espèces peuvent être interprétées comme une adaptation aux conditions respectives de l'habitat, et peuvent être alors un facteur important dans la préférence pour l'habitat et la répartition des deux espèces.

Background: Understanding the factors that shape the distribution of tropical tree species at large scales is a central issue in ecology, conservation and forest management. The aims of this study were to (i) assess the importance of environmental factors relative to historical factors for tree species distributions in the semi-evergreen forests of the northern Congo basin; and to (ii) identify potential mechanisms explaining distribution patterns through a trait-based approach. Methodology/Principal Findings: We analyzed the distribution patterns of 31 common tree species in an area of more than 700,000 km 2 spanning the borders of Cameroon, the Central African Republic, and the Republic of Congo using forest inventory data from 56,445 0.5-ha plots. Spatial variation of environmental (climate, topography and geology) and historical factors (human disturbance) were quantified from maps and satellite records. Four key functional traits (leaf phenology, shade tolerance, wood density, and maximum growth rate) were extracted from the literature. The geological substrate was of major importance for the distribution of the focal species, while climate and past human disturbances had a significant but lesser impact. Species distribution patterns were significantly related to functional traits. Species associated with sandy soils typical of sandstone and alluvium were characterized by slow growth rates, shade tolerance, evergreen leaves, and high wood density, traits allowing persistence on resource-poor soils. In contrast, fast-growing pioneer species rarely occurred on sandy soils, except for Lophira alata. Conclusions/Significance: The results indicate strong environmental filtering due to differential soil resource availability across geological substrates. Additionally, long-term human disturbances in resource-rich areas may have accentuated the observed patterns of species and trait distributions. Trait differences across geological substrates imply pronounced differences in population and ecosystem processes, and call for different conservation and management strategies.