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Understanding the mechanisms that promote the coexistence of hundreds of species over small areas in tropical forest remains a challenge. Many tropical tree species are presumed to be functionally equivalent shade tolerant species but exist on a continuum of performance trade‐offs between survival in shade and the ability to quickly grow in sunlight. These trade‐offs can promote coexistence by reducing fitness differences.Variation in plant functional traits related to resource acquisition is thought to predict variation in performance among species, perhaps explaining community assembly across habitats with gradients in resource availability. Many studies have found low predictive power, however, when linking trait measurements to species demographic rates.Seedlings face different challenges recruiting on the forest floor and may exhibit different traits and/or performance trade‐offs than older individuals face in the eventual adult niche. Seed mass is the typical proxy for seedling success, but species also differ in cotyledon strategy (reserve vs. photosynthetic) or other leaf, stem and root traits. These can cause species with the same average seed mass to have divergent performance in the same habitat.We combined long‐term studies of seedling dynamics with functional trait data collected at a standard life‐history stage in three diverse neotropical forests to ask whether variation in coordinated suites of traits predicts variation among species in demographic performance.Across hundreds of species in Ecuador, Panama and Puerto Rico, we found seedlings displayed correlated suites of leaf, stem, and root traits, which strongly correlated with seed mass and cotyledon strategy. Variation among species in seedling functional traits, seed mass, and cotyledon strategy were strong predictors of trade‐offs in seedling growth and survival. These results underscore the importance of matching the ontogenetic stage of the trait measurement to the stage of demographic dynamics.Our findings highlight the importance of cotyledon strategy in addition to seed mass as a key component of seed and seedling biology in tropical forests because of the contribution of carbon reserves in storage cotyledons to reducing mortality rates and explaining the growth‐survival trade‐off among species.Synthesis: With strikingly consistent patterns across three tropical forests, we find strong evidence for the promise of functional traits to provide mechanistic links between seedling form and demographic performance. [ABSTRACT FROM AUTHOR]

Summary: 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. [ABSTRACT FROM AUTHOR]

Pre‐dispersal seed mortality caused by premature fruit drop is a potentially important source of plant mortality, but one which has rarely been studied in the context of tropical forest plants. Of particular interest is premature fruit drop triggered by enemies, which—if density dependent—could contribute to species coexistence in tropical forest plant communities.We used a long‐term (31 year) dataset on seed and fruit fall obtained through weekly collections from a network of seed traps in a lowland tropical forest (Barro Colorado Island, Panama) to estimate the proportion of seeds prematurely abscised for 201 woody plant species. To determine whether enemy attack might contribute to premature fruit drop, we tested whether plant species abscise more of their fruit prematurely if they (a) have attributes hypothesised to be associated with high levels of enemy attack and (b) are known to be attacked by one enemy group (insect seed predators). We also tested (c) whether mean rates of premature fruit drop for plant species are phylogenetically conserved.Overall rates of premature fruit drop were high in the plant community. Across all species, 39% of seeds were abscised before completing their development. Rates of premature seed abscission varied considerably among species and could not be explained by phylogeny. Premature seed abscission rates were higher in species which are known to host pre‐dispersal insect seed predators and species with attributes that were hypothesised to make them more susceptible to attack by pre‐dispersal enemies, namely species which (a) have larger seeds, (b) have a greater average height, (c) have temporally predictable fruiting patterns and (d) are more abundant at the study site.Synthesis. Premature fruit drop is likely to be a major source of seed mortality for many plant species on Barro Colorado Island. It is plausible that pre‐dispersal seed enemies, such as insect seed predators, contribute to community‐level patterns of premature fruit drop and have the potential to mediate species coexistence through stabilising negative density dependence. Our study suggests that the role of pre‐dispersal enemies in structuring tropical plant communities should be considered alongside the more commonly studied post‐dispersal seed and seedling enemies. [ABSTRACT FROM AUTHOR]

Nutrient addition experiments indicate that nitrogen and phosphorus limit plant processes in many tropical forests. However, the long‐term consequences for forest structure and species composition remain unexplored. We are positioned to evaluate potential long‐term consequences of nutrient addition in central Panama where we have maintained a factorial nitrogen–phosphorus–potassium fertilization experiment for 21 yr and an independent study quantified the species‐specific nutrient requirements of 550 local tree species. Here, we ask whether nutrients limit reproduction at the species and community levels. We also ask whether species‐specific reproductive responses to nutrient addition are stronger among species associated with naturally fertile soils, which could contribute to a shift in species composition. We quantified species‐level reproductive responses for 38 focal species in the 21st year of the experiment and community‐level reproductive litter production for the first 20 yr. Species‐level reproductive responses to nitrogen and potassium addition were weak, inconsistent across species, and insignificant across the 38 focal species. In contrast, species‐level responses to phosphorus addition were consistently and significantly positive across the 38 focal species but were unrelated to species‐specific phosphorus requirements documented independently for the same species. Community‐level reproductive litter production was unaffected by nutrient addition, possibly because spatial and temporal variation is large. We conclude that phosphorus limits reproduction by trees in our experiment but find no evidence that reproductive responses to phosphorus addition favor species associated with naturally phosphorus‐rich soils. [ABSTRACT FROM AUTHOR]

A Thomson Web of Science search for the topic "leaf age" AND potassium AND tropic* identified just one similar study, which reported a similar strong decline in foliar K with leaf age ([16]). This scientific commentary refers to 'Changes in leaf functional traits with leaf age: When do leaves decrease their photosynthetic capacity in Amazonian forests? Google Scholar Crossref Search ADS WorldCat 6 Kitajima K, Mulkey SS, Wright SJ (1997 a) Decline of photosynthetic capacity with leaf age in relation to leaf longevities for five tropical canopy tree species. The LDO model integrates seasonality in the timing of leaf production and abscission, total leaf area present and the contribution of expanding, mature and old leaves to successfully explain seasonality in gross ecosystem production. [Extracted from the article]

Tropical forest productivity represents an important global carbon sink, but many tropical forests grow on infertile soils. Efficient nutrient cycling by litterfall has long been assumed to maintain tropical tree growth, but there is no direct evidence that the nutrients cycled in litterfall are essential for tropical forest productivity.To test whether nutrient cycling by litterfall maintains tropical forest above‐ground productivity, we established large‐scale long‐term litter removal (L−) and litter addition (L+) treatments in a mature lowland tropical forest. We hypothesised that the removal of nutrients in litter would reduce tree growth, survival and litter production in L− plots. By contrast, the addition of nutrients in litter would enhance tree growth, survival and litter production in L+ plots. To test our hypotheses, we recorded tree growth and survival every 2 years, and measured litterfall monthly during 17 years of treatments.Tree growth and litterfall declined over time in L− plots, with consistently lower growth rates compared to controls after 8 years, and lower litter production after 4 years of treatments. By contrast, although litterfall was higher in the L+ plots relative to the controls, there was only a minor transient increase in tree growth immediately after the start of treatments. Tree survival declined over time in all treatments but was not affected by litter manipulation.The long‐term decline in tree growth and litterfall in the L− plots provides the first empirical evidence that nutrient cycling by litterfall plays a key role in maintaining above‐ground productivity in this tropical forest. By contrast, the transient increase in growth in the L+ plots can be attributed to the large inputs of nutrients with the addition of the entire litter standing crop at the start of treatments. The addition of nutrients in litter over the long term was nonetheless sufficient to enhance litter production, possibly by accelerating leaf turnover.Synthesis: Efficient nutrient cycling by litterfall makes an important contribution to the annual nutrient requirements of mature tropical forest trees, compensating for infertile soils. Disturbances that disrupt this finely balanced cycle could therefore reduce biomass carbon sequestration in tropical forests. [ABSTRACT FROM AUTHOR]

Summary: 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. [ABSTRACT FROM AUTHOR]

Summary: Tropical forest root characteristics and resource acquisition strategies are underrepresented in vegetation and global models, hampering the prediction of forest–climate feedbacks for these carbon‐rich ecosystems. Lowland tropical forests often have globally unique combinations of high taxonomic and functional biodiversity, rainfall seasonality, and strongly weathered infertile soils, giving rise to distinct patterns in root traits and functions compared with higher latitude ecosystems. We provide a roadmap for integrating recent advances in our understanding of tropical forest belowground function into vegetation models, focusing on water and nutrient acquisition. We offer comparisons of recent advances in empirical and model understanding of root characteristics that represent important functional processes in tropical forests. We focus on: (1) fine‐root strategies for soil resource exploration, (2) coupling and trade‐offs in fine‐root water vs nutrient acquisition, and (3) aboveground–belowground linkages in plant resource acquisition and use. We suggest avenues for representing these extremely diverse plant communities in computationally manageable and ecologically meaningful groups in models for linked aboveground–belowground hydro‐nutrient functions. Tropical forests are undergoing warming, shifting rainfall regimes, and exacerbation of soil nutrient scarcity caused by elevated atmospheric CO2. The accurate model representation of tropical forest functions is crucial for understanding the interactions of this biome with the climate. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Summary: 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 CO2 and water fluxes.Comparison between observed and TBM trait gradients showed divergence that impacted canopy‐scale simulations of water vapor and CO2 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. [ABSTRACT FROM AUTHOR]

Summary: The representation of stomatal regulation of transpiration and CO2 assimilation is key to forecasting terrestrial ecosystem responses to global change. Given its importance in determining the relationship between forest productivity and climate, accurate and mechanistic model representation of the relationship between stomatal conductance (gs) and assimilation is crucial.We assess possible physiological and mechanistic controls on the estimation of the g1 (stomatal slope, inversely proportional to water use efficiency) and g0 (stomatal intercept) parameters, using diurnal gas exchange surveys and leaf‐level response curves of six tropical broadleaf evergreen tree species.g1 estimated from ex situ response curves averaged 50% less than g1 estimated from survey data. While g0 and g1 varied between leaves of different phenological stages, the trend was not consistent among species. We identified a diurnal trend associated with g1 and g0 that significantly improved model projections of diurnal trends in transpiration.The accuracy of modeled gs can be improved by accounting for variation in stomatal behavior across diurnal periods, and between measurement approaches, rather than focusing on phenological variation in stomatal behavior. Additional investigation into the primary mechanisms responsible for diurnal variation in g1 will be required to account for this phenomenon in land‐surface models. [ABSTRACT FROM AUTHOR]

Topographic heterogeneity and lateral subsurface flow at the hillslope scale of ≤1 km may have outsized impacts on tropical forest through their impacts on water available to plants under water-stressed conditions. However, vegetation dynamics and finer-scale hydrologic processes are not concurrently represented in Earth system models. In this study, we integrate the Energy Exascale Earth System Model (E3SM) land model (ELM) that includes the Functionally Assembled Terrestrial Ecosystem Simulator (FATES), with a three-dimensional hydrology model (ParFlow) to explicitly resolve hillslope topography and subsurface flow and perform numerical experiments to understand how hillslope-scale hydrologic processes modulate vegetation along water availability gradients at Barro Colorado Island (BCI), Panama. Our simulations show that groundwater table depth (WTD) can play a large role in governing aboveground biomass (AGB) when drought-induced tree mortality is triggered by hydraulic failure. Analyzing the simulations using random forest (RF) models, we find that the domain-wide simulated AGB and WTD can be well predicted by static topographic attributes, including surface elevation, slope, and convexity, and adding soil moisture or groundwater table depth as predictors further improves the RF models. Different model representations of mortality due to hydraulic failure can change the dominant topographic driver for the simulated AGB. Contrary to the simulations, the observed AGB in the well-drained 50 ha forest census plot within BCI cannot be well predicted by the RF models using topographic attributes and observed soil moisture as predictors, suggesting other factors such as nutrient status may have a larger influence on the observed AGB. The new coupled model may be useful for understanding the diverse impact of local heterogeneity by isolating the water availability and nutrient availability from the other external and internal factors in ecosystem modeling. [ABSTRACT FROM AUTHOR]