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Nutrient limitation may constrain the ability of recovering and mature tropical forests to serve as a carbon sink. However, it is unclear to what extent trees can utilize nutrient acquisition strategies - especially root phosphatase enzymes and mycorrhizal symbioses - to overcome low nutrient availability across secondary succession. Using a large-scale, full factorial nitrogen and phosphorus fertilization experiment of 76 plots along a secondary successional gradient in lowland wet tropical forests of Panama, we tested the extent to which root phosphatase enzyme activity and mycorrhizal colonization are flexible, and if investment shifts over succession, reflective of changing nutrient limitation. We also conducted a meta-analysis to test how tropical trees adjust these strategies in response to nutrient additions and across succession. We find that tropical trees are dynamic, adjusting investment in strategies - particularly root phosphatase - in response to changing nutrient conditions through succession. These changes reflect a shift from strong nitrogen to weak phosphorus limitation over succession. Our meta-analysis findings were consistent with our field study; we found more predictable responses of root phosphatase than mycorrhizal colonization to nutrient availability. Our findings suggest that nutrient acquisition strategies respond to nutrient availability and demand in tropical forests, likely critical for alleviating nutrient limitation., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

Climate models predict that everwet western Amazonian forests will face warmer and wetter atmospheric conditions, and increased cloud cover. It remains unclear how these changes will impact plant reproductive performance, such as flowering, which plays a central role in sustaining food webs and forest regeneration. Warmer and wetter nights may cause reduced flower production, via increased dark respiration rates or alteration in the reliability of flowering cue-based processes. Additionally, more persistent cloud cover should reduce the amounts of solar irradiance, which could limit flower production. We tested whether interannual variation in flower production has changed in response to fluctuations in irradiance, rainfall, temperature, and relative humidity over 18 yrs in an everwet forest in Ecuador. Analyses of 184 plant species showed that flower production declined as nighttime temperature and relative humidity increased, suggesting that warmer nights and greater atmospheric water saturation negatively impacted reproduction. Species varied in their flowering responses to climatic variables but this variation was not explained by life form or phylogeny. Our results shed light on how plant communities will respond to climatic changes in this everwet region, in which the impacts of these changes have been poorly studied compared with more seasonal Neotropical areas., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

Growing evidence suggests that liana competition with trees is threatening the global carbon sink by slowing the recovery of forests following disturbance. A recent theory based on local and regional evidence further proposes that the competitive success of lianas over trees is driven by interactions between forest disturbance and climate. We present the first global assessment of liana-tree relative performance in response to forest disturbance and climate drivers. Using an unprecedented dataset, we analysed 651 vegetation samples representing 26,538 lianas and 82,802 trees from 556 unique locations worldwide, derived from 83 publications. Results show that lianas perform better relative to trees (increasing liana-to-tree ratio) when forests are disturbed, under warmer temperatures and lower precipitation and towards the tropical lowlands. We also found that lianas can be a critical factor hindering forest recovery in disturbed forests experiencing liana-favourable climates, as chronosequence data show that high competitive success of lianas over trees can persist for decades following disturbances, especially when the annual mean temperature exceeds 27.8°C, precipitation is less than 1614 mm and climatic water deficit is more than 829 mm. These findings reveal that degraded tropical forests with environmental conditions favouring lianas are disproportionately more vulnerable to liana dominance and thus can potentially stall succession, with important implications for the global carbon sink, and hence should be the highest priority to consider for restoration management., (© 2024 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Flowering and fruiting phenology have been infrequently studied in the ever-wet hyperdiverse lowland forests of northwestern equatorial Amazonía. These Neotropical forests are typically called aseasonal with reference to climate because they are ever-wet, and it is often assumed they are also aseasonal with respect to phenology. The physiological limits to plant reproduction imposed by water and light availability are difficult to disentangle in seasonal forests because these variables are often temporally correlated, and both are rarely studied together, challenging our understanding of their relative importance as drivers of reproduction. Here we report on the first long-term study (18 years) of flowering and fruiting phenology in a diverse equatorial forest, Yasuní in eastern Ecuador, and the first to include a full suite of on-site monthly climate data. Using twice monthly censuses of 200 traps and >1000 species, we determined whether reproduction at Yasuní is seasonal at the community and species levels and analyzed the relationships between environmental variables and phenology. We also tested the hypothesis that seasonality in phenology, if present, is driven primarily by irradiance. Both the community- and species-level measures demonstrated strong reproductive seasonality at Yasuní. Flowering peaked in September-November and fruiting peaked in March-April, with a strong annual signal for both phenophases. Irradiance and rainfall were also highly seasonal, even though no month on average experienced drought (a month with <100 mm rainfall). Flowering was positively correlated with current or near-current irradiance, supporting our hypothesis that the extra energy available during the period of peak irradiance drives the seasonality of flowering at Yasuní. As Yasuní is representative of lowland ever-wet equatorial forests of northwestern Amazonía, we expect that reproductive phenology will be strongly seasonal throughout this region., (© 2023 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

For species to coexist, performance must decline as the density of conspecific individuals increases. Although evidence for such conspecific negative density dependence (CNDD) exists in forests, the within-species spatial repulsion it should produce has rarely been demonstrated in adults. In this study, we show that in comparison to a null model of stochastic birth, death, and limited dispersal, the adults of dozens of tropical forest tree species show strong spatial repulsion, some to surprising distances of approximately 100 meters. We used simulations to show that such strong repulsion can only occur if CNDD considerably exceeds heterospecific negative density dependence-an even stronger condition required for coexistence-and that large-scale repulsion can indeed result from small-scale CNDD. These results demonstrate substantial niche differences between species that may stabilize species diversity.

The benefits of masting (volatile, quasi-synchronous seed production at lagged intervals) include satiation of seed predators, but these benefits come with a cost to mutualist pollen and seed dispersers. If the evolution of masting represents a balance between these benefits and costs, we expect mast avoidance in species that are heavily reliant on mutualist dispersers. These effects play out in the context of variable climate and site fertility among species that vary widely in nutrient demand. Meta-analyses of published data have focused on variation at the population scale, thus omitting periodicity within trees and synchronicity between trees. From raw data on 12 million tree-years worldwide, we quantified three components of masting that have not previously been analysed together: (i) volatility, defined as the frequency-weighted year-to-year variation; (ii) periodicity, representing the lag between high-seed years; and (iii) synchronicity, indicating the tree-to-tree correlation. Results show that mast avoidance (low volatility and low synchronicity) by species dependent on mutualist dispersers explains more variation than any other effect. Nutrient-demanding species have low volatility, and species that are most common on nutrient-rich and warm/wet sites exhibit short periods. The prevalence of masting in cold/dry sites coincides with climatic conditions where dependence on vertebrate dispersers is less common than in the wet tropics. Mutualist dispersers neutralize the benefits of masting for predator satiation, further balancing the effects of climate, site fertility and nutrient demands., (© 2023. The Author(s), under exclusive licence to Springer Nature Limited.)

Terrestrial biosphere models (TBMs) include the representation of vertical gradients in leaf traits associated with modeling photosynthesis, respiration, and stomatal conductance. However, model assumptions associated with these gradients have not been tested in complex tropical forest canopies. We compared TBM representation of the vertical gradients of key leaf traits with measurements made in a tropical forest in Panama and then quantified the impact of the observed gradients on simulated canopy-scale CO 2 and water fluxes. Comparison between observed and TBM trait gradients showed divergence that impacted canopy-scale simulations of water vapor and CO 2 exchange. Notably, the ratio between the dark respiration rate and the maximum carboxylation rate was lower near the ground than at the top-of-canopy, leaf-level water-use efficiency was markedly higher at the top-of-canopy, and the decrease in maximum carboxylation rate from the top-of-canopy to the ground was less than TBM assumptions. The representation of the gradients of leaf traits in TBMs is typically derived from measurements made within-individual plants, or, for some traits, assumed constant due to a lack of experimental data. Our work shows that these assumptions are not representative of the trait gradients observed in species-rich, complex tropical forests., (© 2023 The Authors New Phytologist © 2023 New Phytologist Foundation This article has been contributed to by U.S. Government employees and their work is in the public domain in the USA.)

Tropical tree diversity increases with rainfall 1,2 . Direct physiological effects of moisture availability and indirect effects mediated by biotic interactions are hypothesized to contribute to this pantropical increase in diversity with rainfall 2-6 . Previous studies have demonstrated direct physiological effects of variation in moisture availability on tree survival and diversity 5,7-10 , but the indirect effects of variation in moisture availability on diversity mediated by biotic interactions have not been shown 11 . Here we evaluate the relationships between interannual variation in moisture availability, the strength of density-dependent interactions, and seedling diversity in central Panama. Diversity increased with soil moisture over the first year of life across 20 annual cohorts. These first-year changes in diversity persisted for at least 15 years. Differential survival of moisture-sensitive species did not contribute to the observed changes in diversity. Rather, negative density-dependent interactions among conspecifics were stronger and increased diversity in wetter years. This suggests that moisture availability enhances diversity indirectly through moisture-sensitive, density-dependent conspecific interactions. Pathogens and phytophagous insects mediate interactions among seedlings in tropical forests 12-18 , and many of these plant enemies are themselves moisture-sensitive 19-27 . Changes in moisture availability caused by climate change and habitat degradation may alter these interactions and tropical tree diversity., (© 2023. The Author(s), under exclusive licence to Springer Nature Limited.)

Within vascular plants, the partitioning of hydraulic resistance along the soil-to-leaf continuum affects transpiration and its response to environmental conditions. In trees, the fractional contribution of leaf hydraulic resistance (R leaf ) to total soil-to-leaf hydraulic resistance (R total ), or fR leaf (=R leaf /R total ), is thought to be large, but this has not been tested comprehensively. We compiled a multibiome data set of fR leaf using new and previously published measurements of pressure differences within trees in situ. Across 80 samples, fR leaf averaged 0.51 (95% confidence interval [CI] = 0.46-0.57) and it declined with tree height. We also used the allometric relationship between field-based measurements of soil-to-leaf hydraulic conductance and laboratory-based measurements of leaf hydraulic conductance to compute the average fR leaf for 19 tree samples, which was 0.40 (95% CI = 0.29-0.56). The in situ technique produces a more accurate descriptor of fR leaf because it accounts for dynamic leaf hydraulic conductance. Both approaches demonstrate the outsized role of leaves in controlling tree hydrodynamics. A larger fR leaf may help stems from loss of hydraulic conductance. Thus, the decline in fR leaf with tree height would contribute to greater drought vulnerability in taller trees and potentially to their observed disproportionate drought mortality., (© 2022 John Wiley & Sons Ltd.)

Recent observations suggest that the large carbon sink in mature and recovering forests may be strongly limited by nitrogen 1-3 . Nitrogen-fixing trees (fixers) in symbiosis with bacteria provide the main natural source of new nitrogen to tropical forests 3,4 . However, abundances of fixers are tightly constrained 5-7 , highlighting the fundamental unanswered question of what limits new nitrogen entering tropical ecosystems. Here we examine whether herbivory by animals is responsible for limiting symbiotic nitrogen fixation in tropical forests. We evaluate whether nitrogen-fixing trees experience more herbivory than other trees, whether herbivory carries a substantial carbon cost, and whether high herbivory is a result of herbivores targeting the nitrogen-rich leaves of fixers 8,9 . We analysed 1,626 leaves from 350 seedlings of 43 tropical tree species in Panama and found that: (1) although herbivory reduces the growth and survival of all seedlings, nitrogen-fixing trees undergo 26% more herbivory than non-fixers; (2) fixers have 34% higher carbon opportunity costs owing to herbivory than non-fixers, exceeding the metabolic cost of fixing nitrogen; and (3) the high herbivory of fixers is not driven by high leaf nitrogen. Our findings reveal that herbivory may be sufficient to limit tropical symbiotic nitrogen fixation and could constrain its role in alleviating nitrogen limitation on the tropical carbon sink., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

Vegetation demographic models (VDMs) endeavor to predict how global forests will respond to climate change. This requires simulating which trees, if any, are able to recruit under changing environmental conditions. We present a new recruitment scheme for VDMs in which functional-type-specific recruitment rates are sensitive to light, soil moisture and the productivity of reproductive trees. We evaluate the scheme by predicting tree recruitment for four tropical tree functional types under varying meteorology and canopy structure at Barro Colorado Island, Panama. We compare predictions to those of a current VDM, quantitative observations and ecological expectations. We find that the scheme improves the magnitude and rank order of recruitment rates among functional types and captures recruitment limitations in response to variable understory light, soil moisture and precipitation regimes. Our results indicate that adopting this framework will improve VDM capacity to predict functional-type-specific tree recruitment in response to climate change, thereby improving predictions of future forest distribution, composition and function., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

Among tropical forests, lianas are predicted to have a growth advantage over trees during seasonal drought, with substantial implications for tree and forest dynamics. We tested the hypotheses that lianas maintain higher water status than trees during seasonal drought and that lianas maximize leaf cover to match high, dry-season light conditions, while trees are more limited by moisture availability during the dry season. We monitored the seasonal dynamics of predawn and midday leaf water potentials and leaf phenology for branches of 16 liana and 16 tree species in the canopies of two lowland tropical forests with contrasting rainfall regimes in Panama. In a wet, weakly seasonal forest, lianas maintained higher water balance than trees and maximized their leaf cover during dry-season conditions, when light availability was high, while trees experienced drought stress. In a drier, strongly seasonal forest, lianas and trees displayed similar dry season reductions in leaf cover following strong decreases in soil water availability. Greater soil moisture availability and a higher capacity to maintain water status allow lianas to maintain the turgor potentials that are critical for plant growth in a wet and weakly seasonal forest but not in a dry and strongly seasonal forest., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

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., (© 2021 John Wiley & Sons Ltd.)

Deep-water access is arguably the most effective, but under-studied, mechanism that plants employ to survive during drought. Vulnerability to embolism and hydraulic safety margins can predict mortality risk at given levels of dehydration, but deep-water access may delay plant dehydration. Here, we tested the role of deep-water access in enabling survival within a diverse tropical forest community in Panama using a novel data-model approach. We inversely estimated the effective rooting depth (ERD, as the average depth of water extraction), for 29 canopy species by linking diameter growth dynamics (1990-2015) to vapor pressure deficit, water potentials in the whole-soil column, and leaf hydraulic vulnerability curves. We validated ERD estimates against existing isotopic data of potential water-access depths. Across species, deeper ERD was associated with higher maximum stem hydraulic conductivity, greater vulnerability to xylem embolism, narrower safety margins, and lower mortality rates during extreme droughts over 35 years (1981-2015) among evergreen species. Species exposure to water stress declined with deeper ERD indicating that trees compensate for water stress-related mortality risk through deep-water access. The role of deep-water access in mitigating mortality of hydraulically-vulnerable trees has important implications for our predictive understanding of forest dynamics under current and future climates., (No claim to original US Government works New Phytologist © 2021 New Phytologist Foundation.)

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., Competing Interests: Competing interest statement: C.J.C. and J.C. are coauthors on a 2020 article; S.J.W. and J.C. are coauthors on a 2019 article; J.R.P. and J.C. are coauthors on a 2020 article; M.U., J.K.Z., and J.C. are coauthors on two 2019 articles; and M. Silman, W.F.R., and J.C. are coauthors on 2018, 2019, and 2020 articles.

The effects of climate change on tropical forests will depend on how diverse tropical tree species respond to drought. Current distributions of evergreen and deciduous tree species across local and regional moisture gradients reflect their ability to tolerate drought stress, and might be explained by functional traits. We measured leaf water potential at turgor loss (i.e. 'wilting point'; π tlp ), wood density (WD) and leaf mass per area (LMA) on 50 of the most abundant tree species in central Panama. We then tested their ability to explain distributions of evergreen and deciduous species within a 50 ha plot on Barro Colorado Island and across a 70 km rainfall gradient spanning the Isthmus of Panama. Among evergreen trees, species with lower π tlp were associated with drier habitats, with π tlp explaining 28% and 32% of habitat association on local and regional scales, respectively, greatly exceeding the predictive power of WD and LMA. In contrast, π tlp did not predict habitat associations among deciduous species. Across spatial scales, π tlp is a useful indicator of habitat preference for tropical tree species that retain their leaves during periods of water stress, and holds the potential to predict vegetation responses to climate change., (© 2021 The Authors. New Phytologist © 2021 New Phytologist Foundation.)

Nonstructural carbohydrates (NSCs) are essential for maintenance of plant metabolism and may be sensitive to short- and long-term climatic variation. NSC variation in moist tropical forests has rarely been studied, so regulation of NSCs in these systems is poorly understood. We measured foliar and branch NSC content in 23 tree species at three sites located across a large precipitation gradient in Panama during the 2015-2016 El Niño to examine how short- and long-term climatic variation impact carbohydrate dynamics. There was no significant difference in total NSCs as the drought progressed (leaf P = 0.32, branch P = 0.30) nor across the rainfall gradient (leaf P = 0.91, branch P = 0.96). Foliar soluble sugars decreased while starch increased over the duration of the dry period, suggesting greater partitioning of NSCs to storage than metabolism or transport as drought progressed. There was a large variation across species at all sites, but total foliar NSCs were positively correlated with leaf mass per area, whereas branch sugars were positively related to leaf temperature and negatively correlated with daily photosynthesis and wood density. The NSC homoeostasis across a wide range of conditions suggests that NSCs are an allocation priority in moist tropical forests., (© 2018 John Wiley & Sons Ltd.)

Tree death drives population dynamics, nutrient cycling, and evolution within plant communities. Mortality variation across species is thought to be influenced by different factors relative to variation within species. The unified model provided here separates mortality rates into growth-dependent and growth-independent hazards. This model creates the opportunity to simultaneously estimate these hazards both across and within species. Moreover, it provides the ability to examine how species traits affect growth-dependent and growth-independent hazards. We derive this unified mortality model using cross-validated Bayesian methods coupled with mortality data collected over three census intervals for 203 tropical rainforest tree species at Barro Colorado Island (BCI), Panama. We found that growth-independent mortality tended to be higher in species with lower wood density, higher light requirements, and smaller maximum diameter at breast height (dbh). Mortality due to marginal carbon budget as measured by near-zero growth rate tended to be higher in species with lower wood density and higher light demand. The total mortality variation attributable to differences among species was large relative to variation explained by these traits, emphasizing that much remains to be understood. This additive hazards model strengthens our capacity to parse and understand individual-level mortality in highly diverse tropical forests and hence to predict its consequences., Competing Interests: The authors declare no conflict of interest.

Drought disproportionately affects larger trees in tropical forests, but implications for forest composition and carbon (C) cycling in relation to dry season intensity remain poorly understood. In order to characterize how C cycling is shaped by tree size and drought adaptations and how these patterns relate to spatial and temporal variation in water deficit, we analyze data from three forest dynamics plots spanning a moisture gradient in Panama that have experienced El Niño droughts. At all sites, aboveground C cycle contributions peaked below 50-cm stem diameter, with stems ≥ 50 cm accounting for on average 59% of live aboveground biomass, 45% of woody productivity and 49% of woody mortality. The dominance of drought-avoidance strategies increased interactively with stem diameter and dry season intensity. Although size-related C cycle contributions did not vary systematically across the moisture gradient under nondrought conditions, woody mortality of larger trees was disproportionately elevated under El Niño drought stress. Thus, large (> 50 cm) stems, which strongly mediate but do not necessarily dominate C cycling, have drought adaptations that compensate for their more challenging hydraulic environment, particularly in drier climates. However, these adaptations do not fully buffer the effects of severe drought, and increased large tree mortality dominates ecosystem-level drought responses., (© 2017 Smithsonian Institute New Phytologist © 2017 New Phytologist Trust.)

Mounting evidence suggests that anthropogenic global change is altering plant species composition in tropical forests. Fewer studies, however, have focused on long-term trends in reproductive activity, in part because of the lack of data from tropical sites. Here, we analyze a 28-year record of tropical flower phenology in response to anthropogenic climate and atmospheric change. We show that a multidecadal increase in flower activity is most strongly associated with rising atmospheric CO 2 concentrations using yearly aggregated data. Compared to significant climatic factors, CO 2 had on average an approximately three-, four-, or fivefold stronger effect than rainfall, solar radiation, and the Multivariate ENSO Index, respectively. Peaks in flower activity were associated with greater solar radiation and lower rainfall during El Niño years. The effect of atmospheric CO 2 on flowering has diminished over the most recent decade for lianas and canopy trees, whereas flowering of midstory trees and shrub species continued to increase with rising CO 2 . Increases in flowering were accompanied by a lengthening of flowering duration for canopy and midstory trees. Understory treelets did not show increases in flowering but did show increases in duration. Given that atmospheric CO 2 will likely continue to climb over the next century, a long-term increase in flowering activity may persist in some growth forms until checked by nutrient limitation or by climate change through rising temperatures, increasing drought frequency and/or increasing cloudiness and reduced insolation., (© 2017 John Wiley & Sons Ltd.)

The tropical forests of Borneo and Amazonia may each contain more tree species diversity in half a square kilometre than do all the temperate forests of Europe, North America, and Asia combined. Biologists have long been fascinated by this disparity, using it to investigate potential drivers of biodiversity. Latitudinal variation in many of these drivers is expected to create geographic differences in ecological and evolutionary processes, and evidence increasingly shows that tropical ecosystems have higher rates of diversification, clade origination, and clade dispersal. However, there is currently no evidence to link gradients in ecological processes within communities at a local scale directly to the geographic gradient in biodiversity. Here, we show geographic variation in the storage effect, an ecological mechanism that reduces the potential for competitive exclusion more strongly in the tropics than it does in temperate and boreal zones, decreasing the ratio of interspecific-to-intraspecific competition by 0.25% for each degree of latitude that an ecosystem is located closer to the Equator. Additionally, we find evidence that latitudinal variation in climate underpins these differences; longer growing seasons in the tropics reduce constraints on the seasonal timing of reproduction, permitting lower recruitment synchrony between species and thereby enhancing niche partitioning through the storage effect. Our results demonstrate that the strength of the storage effect, and therefore its impact on diversity within communities, varies latitudinally in association with climate. This finding highlights the importance of biotic interactions in shaping geographic diversity patterns, and emphasizes the need to understand the mechanisms underpinning ecological processes in greater detail than has previously been appreciated.

Premise of the Study: Wood density is the top predictor of growth and mortality rates (vital rates) but with modest explanatory power at best. Stronger links to vital rates are expected if wood density is decomposed into its anatomical properties at sapling and adult stages, since saplings and adults differ in wood traits and vital rates. We examined whether anatomical determinants of wood density and strength of the relationship between wood traits and vital rates shift between saplings and adults., Methods: Using wood segments from near pith (sapling) and near bark (adult) for 20 tree species (three adults each) from Barro Colorado Island, Panama, we quantified wood traits. Vital rates for saplings and adults were obtained from an earlier study., Key Results: Anatomical predictors of wood density were similar for sapling and adult wood, with wood density variation largely explained by fiber lumen area and fiber wall fraction. In sapling wood only, growth rates decreased with fiber wall fraction and increased with fiber lumen area, while mortality rates increased with vessel area but decreased with fiber wall fraction and vessel density., Conclusions: Wood traits of sapling trees provide functional insight into the growth-mortality tradeoff. Sapling wood with relatively large fiber lumen area and wide vessels, enabling faster hydraulic transport but less mechanical strength, is associated with fast growth and high mortality. Sapling wood with relatively more fiber wall and many narrow vessels, enabling greater mechanical strength but slower hydraulic transport, is associated with slow growth and low mortality., (© 2017 Botanical Society of America.)

The response of tropical forests to anthropogenic climate change is critically important to future global carbon budgets, yet remains highly uncertain. Here, we investigate how precipitation, temperature, solar radiation and dry- and wet-season lengths are related to annual tree growth, flower production, and fruit production in three moist tropical forest tree species using long-term datasets from tree rings and litter traps in central Panama. We also evaluated how growth, flower, and fruit production were interrelated. We found that growth was positively correlated with wet-season precipitation in all three species: Jacaranda copaia (r = 0.63), Tetragastris panamensis (r = 0.39) and Trichilia tuberculata (r = 0.39). Flowering and fruiting in Jacaranda were negatively related to current-year dry-season rainfall and positively related to prior-year dry-season rainfall. Flowering in Tetragastris was negatively related to current-year annual mean temperature while Trichilia showed no significant relationships of reproduction with climate. Growth was significantly related to reproduction only in Tetragastris, where it was positively related to previous year fruiting. Our results suggest that tree growth in moist tropical forest tree species is generally reduced by drought events such as those associated with strong El Niño events. In contrast, interannual variation in reproduction is not generally associated with growth and has distinct and species-specific climate responses, with positive effects of El Niño events in some species. Understanding these contrasting climate effects on tree growth and reproduction is critical to predicting changes in tropical forest dynamics and species composition under climate change.

Background: Tropical rainforests are frequently limited by soil nutrient availability. However, the response of the metabolic phenotypic plasticity of trees to an increase of soil nutrient availabilities is poorly understood. We expected that increases in the ability of a nutrient that limits some plant processes should be detected by corresponding changes in plant metabolome profile related to such processes., Methodology/principal Findings: We studied the foliar metabolome of saplings of three abundant tree species in a 15 year field NPK fertilization experiment in a Panamanian rainforest. The largest differences were among species and explained 75% of overall metabolome variation. The saplings of the large canopy species, Tetragastris panamensis, had the lowest concentrations of all identified amino acids and the highest concentrations of most identified secondary compounds. The saplings of the "mid canopy" species, Alseis blackiana, had the highest concentrations of amino acids coming from the biosynthesis pathways of glycerate-3P, oxaloacetate and α-ketoglutarate, and the saplings of the low canopy species, Heisteria concinna, had the highest concentrations of amino acids coming from the pyruvate synthesis pathways., Conclusions/significance: The changes in metabolome provided strong evidence that different nutrients limit different species in different ways. With increasing P availability, the two canopy species shifted their metabolome towards larger investment in protection mechanisms, whereas with increasing N availability, the sub-canopy species increased its primary metabolism. The results highlighted the proportional distinct use of different nutrients by different species and the resulting different metabolome profiles in this high diversity community are consistent with the ecological niche theory.

It was recently proposed that boron might be the most important nutrient structuring tree species distributions in tropical forests. Here we combine observational and experimental studies to test this hypothesis for lowland tropical forests of Panama. Plant-available boron is uniformly low in tropical forest soils of Panama and is not significantly associated with any of the > 500 species in a regional network of forest dynamics plots. Experimental manipulation of boron supply to seedlings of three tropical tree species revealed no evidence of boron deficiency or toxicity at concentrations likely to occur in tropical forest soils. Foliar boron did not correlate with soil boron along a local scale gradient of boron availability. Fifteen years of boron addition to a tropical forest increased plant-available boron by 70% but did not significantly change tree productivity or boron concentrations in live leaves, wood or leaf litter. The annual input of boron in rainfall accounts for a considerable proportion of the boron in annual litterfall and is similar to the pool of plant-available boron in the soil, and is therefore sufficient to preclude boron deficiency. We conclude that boron does not influence tree species distributions in Panama and presumably elsewhere in the lowland tropics., (No claim to original US government works New Phytologist © 2016 New Phytologist Trust.)

Dioecy has a demographic disadvantage compared with hermaphroditism: only about half of reproductive adults produce seeds. Dioecious species must therefore have fitness advantages to compensate for this cost through increased survival, growth, and/or reproduction. We used a full life cycle approach to quantify the demographic costs and benefits associated with dioecy while controlling for demographic differences between dioecious and hermaphroditic species related to other functional traits. The advantage of this novel approach is that we can focus on the effect of breeding system across a diverse tree community. We built a composite integral projection model for hermaphroditic and dioecious tree populations from Barro Colorado Island, Panama, using long-term demographic and newly collected reproductive data. Integration of all costs and benefits showed that compensation was realized through increased seed production, resulting in no net costs of dioecy. Compensation was also facilitated by the low contribution of reproduction to population growth. Estimated positive effects of dioecy on tree growth and survival were small and insignificant for population growth rates. Our model revealed that, for long-lived organisms, the cost of having males is smaller than generally expected. Hence, little compensation is required for dioecious species to maintain population growth rates similar to those of hermaphroditic species.

Specialist herbivores and pathogens could induce negative conspecific density dependence among their hosts and thereby contribute to the diversity of plant communities. A small number of hyperdiverse genera comprise a large portion of tree diversity in tropical forests. These closely related congeners are likely to share natural enemies. Diverse defenses could still allow congeners to partition niche space defined by natural enemies, but interspecific differences in defenses would have to exceed intraspecific variation in defenses. We ask whether interspecific variation in secondary chemistry exceeds intraspecific variation for species from four hyperdiverse tropical tree genera. We used novel methods to quantify chemical structural similarity for all compounds present in methanol extracts of leaf tissue. We sought to maximize intraspecific variation by selecting conspecific leaves from different ontogenetic stages (expanding immature vs. fully hardened mature), different light environments (deep understory shade vs. large forest gaps), and different seasons (dry vs. wet). Chemical structural similarity differed with ontogeny, light environment, and season, but interspecific differences including those among congeneric species were much larger. Our results suggest that species differences in secondary chemistry are large relative to within-species variation, perhaps sufficiently large to permit niche segregation among congeneric tree species based on chemical defenses., (© 2016 by the Ecological Society of America.)

Wood structure differs widely among tree species and species with faster growth, higher mortality and larger maximum size have been reported to have fewer but larger vessels and higher hydraulic conductivity (Kh). However, previous studies compiled data from various sources, often failed to control tree size and rarely controlled variation in other traits. We measured wood density, tree size and vessel traits for 325 species from a wet forest in Panama, and compared wood and leaf traits to demographic traits using species-level data and phylogenetically independent contrasts. Wood traits showed strong phylogenetic signal whereas pairwise relationships between traits were mostly phylogenetically independent. Trees with larger vessels had a lower fraction of the cross-sectional area occupied by vessel lumina, suggesting that the hydraulic efficiency of large vessels permits trees to dedicate a larger proportion of the wood to functions other than water transport. Vessel traits were more strongly correlated with the size of individual trees than with maximal size of a species. When individual tree size was included in models, Kh scaled positively with maximal size and was the best predictor for both diameter and biomass growth rates, but was unrelated to mortality., (© 2016 The Authors. New Phytologist © 2016 New Phytologist Trust.)

The phyllosphere (comprising the leaf surface and interior) is one of the world's largest microbial habitats and is host to an abundant and diverse array of bacteria. Nonetheless, the degree to which bacterial communities are benign, harmful, or beneficial to plants in situ is unknown. We tested the hypothesis that the net effect of reducing bacterial abundance and diversity would vary substantially among host species (from harmful to beneficial) and this would be strongly mediated by soil resource availability. To test this, we monitored tree seedling growth responses to commercial antibiotics among replicated resource supply treatments (N, P, K) in a tropical forest in Panama for 29 months. We applied either antibiotics or control water to replicated seedlings of five common tree species (Alseis blackiana, Desmopsis panamensis, Heisteria concinna, Sorocea affinis, and Tetragastris panamensis). These antibiotic treatments significantly reduced both the abundance and diversity of bacteria epiphytically as well as endophytically. Overall, the effect of antibiotics on performance was highly host specific. Applying antibiotics increased growth for three species by as much as 49% (Alseis, Heisteria, and Tetragastris), decreased growth for a fourth species by nearly 20% (Sorocea), and had no impact on a fifth species (Desmopsis). Perhaps more importantly, the degree to which foliar bacteria were harmful or not varied with soil resource supply. Specifically, applying antibiotics had no effect when potassium was added but increased growth rate by almost 40% in the absence of potassium. Alternatively, phosphorus enrichment caused the effect of bacteria to switch from being primarily beneficial to harmful or vice versa, but this depended entirely on the presence or absence of nitrogen enrichment (i.e., important and significant interactions). Our results are the first to demonstrate that the net effect of reducing the abundance and diversity of bacteria can have very strong positive and negative effects on seedling performance. Moreover, these effects were clearly mediated by soil resource availability. Though speculative, we suggest that foliar bacteria may interact with soil fertility to comprise an important, yet cryptic dimension of niche differentiation, which can have important implications for species coexistence., (© 2016 by the Ecological Society of America.)

As distinct community assembly processes can produce similar community patterns, assessing the ecological mechanisms promoting coexistence in hyperdiverse rainforests remains a considerable challenge. We use spatially explicit neighbourhood models of tree growth to quantify how functional trait and phylogenetic similarities predict variation in growth and crowding effects for the 315 most abundant tree species in a 25-ha lowland rainforest plot in Ecuador. We find that functional trait differences reflect variation in (1) species maximum potential growth, (2) the intensity of interspecific interactions for some species, and (3) species sensitivity to neighbours. We find that neighbours influenced tree growth in 28% of the 315 focal tree species. Neighbourhood effects are not detected in the remaining 72%, which may reflect the low statistical power to model rare taxa and/or species insensitivity to neighbours. Our results highlight the spectrum of ways in which functional trait differences can shape community dynamics in highly diverse rainforests., (© 2016 John Wiley & Sons Ltd/CNRS.)

Multiple niche-based processes including conspecific negative density dependence (CNDD) determine plant regeneration and community structure. We ask how interspecific and intraspecific density-dependent interactions relate to plant life histories and associated functional traits. Using hierarchical models, we analysed how such interactions affected first-year survival of seedling recruits of 175 species in a tropical forest, and how species abundances and functional traits are related to interspecific variation in density-dependent effects. Conspecific seedling neighbour effects prevailed over the effects of larger conspecific and all heterospecific neighbours. Tolerance of seedling CNDD enhanced recruit survival and subsequent abundance, all of which were greater among larger seeded, slow-growing and well-defended species. Niche differentiation along the growth-survival trade-off and tolerance of seedling CNDD strongly correlated with regeneration success, with manifest consequences for community structure. The ability of larger seeded species to better tolerate CNDD suggests a novel mechanism for CNDD to contribute to seed-size variation and promote species coexistence through a tolerance-fecundity trade-off., (© 2016 John Wiley & Sons Ltd/CNRS.)

Numerous grassland experiments have found evidence for a complementarity effect, an increase in productivity with higher plant species richness due to niche partitioning. However, empirical tests of complementarity in natural forests are rare. We conducted a spatially explicit analysis of 518 433 growth records for 274 species from a 50-ha tropical forest plot to test neighborhood complementarity, the idea that a tree grows faster when it is surrounded by more dissimilar neighbors. We found evidence for complementarity: focal tree growth rates increased by 39.8% and 34.2% with a doubling of neighborhood multi-trait dissimilarity and phylogenetic dissimilarity, respectively. Dissimilarity from neighbors in maximum height had the most important effect on tree growth among the six traits examined, and indeed, its effect trended much larger than that of the multitrait dissimilarity index. Neighborhood complementarity effects were strongest for light-demanding species, and decreased in importance with increasing shade tolerance of the focal individuals. Simulations demonstrated that the observed neighborhood complementarities were sufficient to produce positive stand-level biodiversity-productivity relationships. We conclude that neighborhood complementarity is important for productivity in this tropical forest, and that scaling down to individual-level processes can advance our understanding of the mechanisms underlying stand-level biodiversity-productivity relationships.

Phenotypic traits and their associated trade-offs have been shown to have globally consistent effects on individual plant physiological functions, but how these effects scale up to influence competition, a key driver of community assembly in terrestrial vegetation, has remained unclear. Here we use growth data from more than 3 million trees in over 140,000 plots across the world to show how three key functional traits--wood density, specific leaf area and maximum height--consistently influence competitive interactions. Fast maximum growth of a species was correlated negatively with its wood density in all biomes, and positively with its specific leaf area in most biomes. Low wood density was also correlated with a low ability to tolerate competition and a low competitive effect on neighbours, while high specific leaf area was correlated with a low competitive effect. Thus, traits generate trade-offs between performance with competition versus performance without competition, a fundamental ingredient in the classical hypothesis that the coexistence of plant species is enabled via differentiation in their successional strategies. Competition within species was stronger than between species, but an increase in trait dissimilarity between species had little influence in weakening competition. No benefit of dissimilarity was detected for specific leaf area or wood density, and only a weak benefit for maximum height. Our trait-based approach to modelling competition makes generalization possible across the forest ecosystems of the world and their highly diverse species composition.

Remote identification and mapping of canopy tree species can contribute valuable information towards our understanding of ecosystem biodiversity and function over large spatial scales. However, the extreme challenges posed by highly diverse, closed-canopy tropical forests have prevented automated remote species mapping of non-flowering tree crowns in these ecosystems. We set out to identify individuals of three focal canopy tree species amongst a diverse background of tree and liana species on Barro Colorado Island, Panama, using airborne imaging spectroscopy data. First, we compared two leading single-class classification methods--binary support vector machine (SVM) and biased SVM--for their performance in identifying pixels of a single focal species. From this comparison we determined that biased SVM was more precise and created a multi-species classification model by combining the three biased SVM models. This model was applied to the imagery to identify pixels belonging to the three focal species and the prediction results were then processed to create a map of focal species crown objects. Crown-level cross-validation of the training data indicated that the multi-species classification model had pixel-level producer's accuracies of 94-97% for the three focal species, and field validation of the predicted crown objects indicated that these had user's accuracies of 94-100%. Our results demonstrate the ability of high spatial and spectral resolution remote sensing to accurately detect non-flowering crowns of focal species within a diverse tropical forest. We attribute the success of our model to recent classification and mapping techniques adapted to species detection in diverse closed-canopy forests, which can pave the way for remote species mapping in a wider variety of ecosystems.

Unlabelled: •, Premise of the Study: Wood specific gravity (WSG) mediates an interspecific trade-off between growth and mortality and is a key measure for estimating carbon stocks. Most studies use species mean values to represent WSG, despite variation at different levels of biological organization. We examined sources of variation in WSG across four nested scales (segments within cores, cores within trees, trees within species, and species), compared the pattern of radial variation in WSG among species differing in growth strategies, and investigated the effect of WSG radial variation on aboveground biomass estimates.•, Methods: We took two perpendicular cores from six individuals each of 20 tropical tree species representing a broad range of mean WSGs and growth-mortality strategies in a lowland tropical moist forest in Panama. Cores were divided into 1-cm segments, and WSG was determined for each segment.•, Key Results: The bulk of the total variance in WSG was dominated by interspecies variation (88%), whereas variation due to measurement error, segments within cores, and cores within trees (8%) was minimal. Radial variation in WSG, defined as change in WSG with increasing distance from the pith, was significant in 17 of the 20 species and included significant monotonic increases in 6 species and nonmonotonic patterns in 11 species. Radial variation in WSG resulted in a small but significant bias in aboveground biomass estimates.•, Conclusions: Radial variation in WSG is related to a species' growth strategy and, though minimal compared with interspecific variation in WSG, can cause a downward bias when not incorporated into aboveground biomass estimates., (© 2014 Botanical Society of America, Inc.)

A complex set of interactions among neighbors influences plant performance and community structure. Understanding their joint operation requires extensive information on species characteristics and individual performance. We evaluated first-year survival of 35719 tropical forest seedlings of 222 species and 15 annual cohorts relative to the density of conspecific and heterospecific neighbors and the phylogenetic similarity of heterospecific neighbors. Neighbors were from two size classes, and size asymmetric interactions provided insight into likely mechanisms. Large heterospecific and conspecific neighbors reduced seedling survival equally, suggesting resource competition rather than host-specific enemies as a mechanism. In contrast, much stronger negative conspecific effects were associated with seedling neighbors capable of limited resource uptake, suggesting shared pests rather than competition as the mechanism. Survival improved, however, near phylogenetically similar heterospecific neighbors, suggesting habitat associations shared among closely related species affect spatial patterns of performance. Improved performance near phylogenetically similar neighbors is an emerging pattern in the handful of similar studies.

Background: Many tropical forest tree species delay greening their leaves until full expansion. This strategy is thought to provide newly flushing leaves with protection against damage by herbivores by keeping young leaves devoid of nutritive value. Because young leaves suffer the greatest predation from invertebrate herbivores, delayed greening could prevent costly tissue loss. Many species that delay greening also produce anthocyanin pigments in their new leaves, giving them a reddish tint. These anthocyanins may be fungicidal, protect leaves against UV damage or make leaves cryptic to herbivores blind to the red part of the spectrum., Methods: A comprehensive survey was undertaken of seedlings, saplings and mature trees in two diverse tropical forests: a rain forest in western Amazonia (Yasuní National Park, Ecuador) and a deciduous forest in Central America (Barro Colorado Island, Panamá). A test was made of whether individuals and species with delayed greening or red-coloured young leaves showed lower mortality or higher relative growth rates than species that did not., Key Results: At both Yasuní and Barro Colorado Island, species with delayed greening or red young leaves comprised significant proportions of the seedling and tree communities. At both sites, significantly lower mortality was found in seedlings and trees with delayed greening and red-coloured young leaves. While there was little effect of leaf colour on the production of new leaves of seedlings, diameter relative growth rates of small trees were lower in species with delayed greening and red-coloured young leaves than in species with regular green leaves, and this effect remained when the trade-off between mortality and growth was accounted for., Conclusions: Herbivores exert strong selection pressure on seedlings for the expression of defence traits. A delayed greening or red-coloured young leaf strategy in seedlings appears to be associated with higher survival for a given growth rate, and may thus influence the species composition of later life stages.

Background and Aims: Leaf life span is widely recognized as a key life history trait associated with herbivory resistance, but rigorous comparative data are rare for seedlings. The goal of this study was to examine how light environment affects leaf life span, and how ontogenetic development during the first year may influence leaf fracture toughness, lamina density and stem density that are relevant for herbivory resistance, leaf life span and seedling survival., Methods: Data from three experiments encompassing 104 neotropical woody species were combined. Leaf life span, lamina and vein fracture toughness, leaf and stem tissue density and seedling survival were quantified for the first-year seedlings at standardized ontogenetic stages in shade houses and common gardens established in gaps and shaded understorey in a moist tropical forest in Panama. Mortality of naturally recruited seedlings till 1 year later was quantified in 800 1-m² plots from 1994 to 2011., Key Results: Median leaf life span ranged widely among species, always greater in shade (ranging from 151 to >1790 d in the understorey and shade houses) than in gaps (115-867 d), but with strong correlation between gaps and shade. Leaf and stem tissue density increased with seedling age, whereas leaf fracture toughness showed only a weak increase. All these traits were positively correlated with leaf life span. Leaf life span and stem density were negatively correlated with seedling mortality in shade, while gap mortality showed no correlation with these traits., Conclusions: The wide spectrum of leaf life span and associated functional traits reflects variation in shade tolerance of first-year seedlings among coexisting trees, shrubs and lianas in this neotropical forest. High leaf tissue density is important in enhancing leaf toughness, a known physical defence, and leaf life span. Both seedling leaf life span and stem density should be considered as key functional traits that contribute to seedling survival in tropical forest understoreys.

Tree deaths open gaps in closed-canopy forests, which allow light to reach the forest floor and promote seed germination and seedling establishment. Gap dependence of regeneration is an important axis of life history variation among forest plant species, and many studies have evaluated how plant species differ in seedling and sapling performance in gaps. However, relatively little is known about how seed arrival in gaps compares with seed arrival in the understory, even though seed dispersal by wind and animals is expected to be altered in gaps. We documented seed arrival for the first seven years after gap formation in the moist tropical forests of Barro Colorado Island (BCI), Panama, and evaluated how the amount and functional composition of arriving seeds compared with understory sites. On average, in the first three years after gap formation, 72% fewer seeds arrived in gaps than in the understory (207 vs. 740 seeds x m(-2) x yr(-1)). The reduction in number of arriving seeds fell disproportionately on animal-dispersed species, which suffered an 86% reduction in total seed number, while wind-dispersed species experienced only a 47% reduction, and explosively dispersed species showed increased seed numbers arriving. The increase in explosively dispersed seeds consisted entirely of the seeds of several shrub species, a result consistent with greater in situ seed production by explosively dispersed shrubs that survived gap formation or recruited immediately thereafter. Lianas did relatively better in seed arrival into gaps than did trees, suffering less of a reduction in seed arrival compared with understory sites. This result could in large part be explained by the greater predominance of wind dispersal among lianas: there were no significant differences between lianas and trees when controlling for dispersal syndromes. Our results show that seed arrival in gaps is very different from seed arrival in the understory in both total seeds arriving and functional composition. Differential seed arrival in gaps will help to maintain wind-dispersed, explosively dispersed, and possibly other understory species in the community of plants that regenerate in gaps.

Leaf dark respiration (R) and its temperature sensitivity are essential for efforts to model carbon fluxes in tropical forests under current and future temperature regimes, but insufficient data exist to generalize patterns of R in species-rich tropical forests. Here, we tested the hypothesis that R and its temperature sensitivity (expressed as Q10, the proportional increase in R with a 10 °C rise in temperature) vary in relation to leaf functional traits, and among plant functional types (PFTs). We conducted in situ measurements of R of 461 leaves of 26 species of tree and liana in the upper canopy of a tropical forest in Panama. A construction crane allowed repeated non-destructive access to measure leaves kept in the dark since the previous night and equilibrated to the ambient temperature of 23-31 °C in the morning. R at 25 °C (R25) varied among species (mean 1.11 μmol m(-2) s(-1); range 0.72-1.79 μmol m(-2) s(-1)) but did not differ significantly among PFTs. R25 correlated positively with photosynthetic capacity, leaf mass per unit area, concentrations of nitrogen and phosphorus, and negatively with leaf lifespan. Q10 estimated for each species was on average higher than the 2.0 often assumed in coupled climate-vegetation models (mean 2.19; range 1.24-3.66). Early-successional tree species had higher Q10 values than other functional types, but interspecific variation in Q10 values was not correlated with other leaf traits. Similarity in respiration characteristics across PFTs, and relatively strong correlations of R with other leaf functional traits offer potential for trait-based vegetation modeling in species-rich tropical forests.

Relationships between functional traits and average or potential demographic rates have provided insight into the functional constraints and trade-offs underlying life-history strategies of tropical tree species. We have extended this framework by decomposing growth rates of -130 000 trees of 171 Neotropical tree species into intrinsic growth and the response of growth to light and size. We related these growth characteristics to multiple functional traits (wood density, adult stature, seed mass, leaf traits) in a hierarchical Bayesian model that accounted for measurement error and intraspecific variability of functional traits. Wood density was the most important trait determining all three growth characteristics. Intrinsic growth rates were additionally strongly related to adult stature, while all traits contributed to light response. Our analysis yielded a predictive model that allows estimation of growth characteristics for rare species on the basis of a few easily measurable morphological traits.

The rapid disruption of tropical forests probably imperils global biodiversity more than any other contemporary phenomenon. With deforestation advancing quickly, protected areas are increasingly becoming final refuges for threatened species and natural ecosystem processes. However, many protected areas in the tropics are themselves vulnerable to human encroachment and other environmental stresses. As pressures mount, it is vital to know whether existing reserves can sustain their biodiversity. A critical constraint in addressing this question has been that data describing a broad array of biodiversity groups have been unavailable for a sufficiently large and representative sample of reserves. Here we present a uniquely comprehensive data set on changes over the past 20 to 30 years in 31 functional groups of species and 21 potential drivers of environmental change, for 60 protected areas stratified across the world’s major tropical regions. Our analysis reveals great variation in reserve ‘health’: about half of all reserves have been effective or performed passably, but the rest are experiencing an erosion of biodiversity that is often alarmingly widespread taxonomically and functionally. Habitat disruption, hunting and forest-product exploitation were the strongest predictors of declining reserve health. Crucially, environmental changes immediately outside reserves seemed nearly as important as those inside in determining their ecological fate, with changes inside reserves strongly mirroring those occurring around them. These findings suggest that tropical protected areas are often intimately linked ecologically to their surrounding habitats, and that a failure to stem broad-scale loss and degradation of such habitats could sharply increase the likelihood of serious biodiversity declines.

The storage effect is a mechanism that can facilitate the coexistence of competing species through temporal fluctuations in reproductive output. Numerous natural systems have the prerequisites for the storage effect, yet it has rarely been quantitatively assessed. Here, we investigate the possible importance of the storage effect in explaining the coexistence of tree species in the diverse tropical forest on Barro Colorado Island, Panama. This tropical forest has been monitored for more than 20 years, and annual seed production is asynchronous among species, a primary requirement for the storage effect. We constructed a model of forest regeneration that includes species-specific recruitment through seed, sapling, and adult stages, and we parameterized the model using data for 28 species for which information is known about seedling germination and survival. Simulations of the model demonstrated that the storage effect alone can be a strong mechanism allowing long-term persistence of species. We also developed a metric to quantify the strength of the storage effect in a way comparable to classical resource partitioning. Applying this metric to seed production data from 108 species, the storage effect reduces the strength of pairwise interspecific competition to 11-43% of the strength of intraspecific competition, thereby demonstrating strong potential to facilitate coexistence. Finally, for a subset of 51 species whose phylogenetic relationships are known, we compared the strength of the storage effect between pairs of species to their phylogenetic similarity. The strength of the storage effect between closely related species was on average no different from distantly related species, implying that the storage effect can be important in promoting the coexistence of even closely related species.

The degree to which turnover in biological communities is structured by deterministic or stochastic factors and the identities of influential deterministic factors are fundamental, yet unresolved, questions in ecology. Answers to these questions are particularly important for projecting the fate of forests with diverse disturbance histories worldwide. To uncover the processes governing turnover we use species-level molecular phylogenies and functional trait data sets for two long-term tropical forest plots with contrasting disturbance histories: one forest is older-growth, and one was recently disturbed. Having both phylogenetic and functional information further allows us to parse out the deterministic influences of different ecological filters. With the use of null models we find that compositional turnover was random with respect to phylogeny on average, but highly nonrandom with respect to measured functional traits. Furthermore, as predicted by a deterministic assembly process, the older-growth and disturbed forests were characterized by less than and greater than expected functional turnover, respectively. These results suggest that the abiotic environment, which changes due to succession in the disturbed forest, strongly governs the temporal dynamics of disturbed and undisturbed tropical forests. Predicting future changes in the composition of disturbed and undisturbed forests may therefore be tractable when using a functional-trait-based approach.

The distribution of plant species along environmental gradients is expected to be predictable based on organismal function. Plant functional trait research has shown that trait values generally vary predictably along broad-scale climatic and soil gradients. This work has also demonstrated that at any one point along these gradients there is a large amount of interspecific trait variation. The present research proposes that this variation may be explained by the local-scale sorting of traits along soil fertility and acidity axes. Specifically, we predicted that trait values associated with high resource acquisition and growth rates would be found on soils that are more fertile and less acidic. We tested the expected relationships at the species-level and quadrat-level (20 × 20 m) using two large forest plots in Panama and China that contain over 450 species combined. Predicted relationships between leaf area and wood density and soil fertility were supported in some instances, but the majority of the predicted relationships were rejected. Alternative resource axes, such as light gradients, therefore likely play a larger role in determining the interspecific variability in plant functional traits in the two forests studied.

Deposition of reactive nitrogen (N) from human activities has large effects on temperate forests where low natural N availability limits productivity but is not known to affect tropical forests where natural N availability is often much greater. Leaf N and the ratio of N isotopes (δ(15)N) increased substantially in a moist forest in Panama between ~1968 and 2007, as did tree-ring δ(15)N in a dry forest in Thailand over the past century. A decade of fertilization of a nearby Panamanian forest with N caused similar increases in leaf N and δ(15)N. Therefore, our results indicate regional increases in N availability due to anthropogenic N deposition. Atmospheric nitrogen dioxide measurements and increased emissions of anthropogenic reactive N over tropical land areas suggest that these changes are widespread in tropical forests.

We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40 x 40 m each. The addition of K was associated with significant decreases in stand-level fine-root biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1-10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (P = 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles.