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Forecasting insect responses to environmental variables at local and global spatial scales remains a crucial task in Ecology. However, predicting future responses requires long-term datasets, which are rarely available for insects, especially in the tropics. From 2002 to 2017, we recorded male ant incidence of 155 ant species at ten malaise traps on the 50-ha ForestGEO plot in Barro Colorado Island. In this Panamanian tropical rainforest, traps were deployed for two weeks during the wet and dry seasons. Short-term changes in the timing of male flying activity were pronounced, and compositionally distinct assemblages flew during the wet and dry seasons. Notably, the composition of these distinct flying assemblages oscillated in consistent 4-year cycles but did not change during the 16-year study period. Across time, a Seasonal Auto-Regressive Integrated Moving Average model explained 75% of long-term variability in male ant production (i.e., the summed incidence of male species across traps), which responded negatively to monthly maximum temperature, and positively to sea surface temperature, a surrogate for El Niño Southern Oscillation (ENSO) events. Establishing these relationships allowed us to forecast ant production until 2022 when year-long local climate variables were available. Consistent with the data, the forecast indicated no significant changes in long-term temporal trends of male ant production. However, simulations of different scenarios of climate variables found that strong ENSO events and maximum temperature impacted male ant production positively and negatively, respectively. Our results highlight the dependence of ant male production on both short- and long-term temperature changes, which is critical under current global warming., Competing Interests: Declarations. Competing interests: The authors declare no competing interests., (© 2024. The Author(s).)

All species must partition resources among the processes that underly growth, survival, and reproduction. The resulting demographic trade-offs constrain the range of viable life-history strategies and are hypothesized to promote local coexistence. Tropical forests pose ideal systems to study demographic trade-offs as they have a high diversity of coexisting tree species whose life-history strategies tend to align along two orthogonal axes of variation: a growth-survival trade-off that separates species with fast growth from species with high survival and a stature-recruitment trade-off that separates species that achieve large stature from species with high recruitment. As these trade-offs have typically been explored for trees ≥1 cm dbh, it is unclear how species' growth and survival during earliest seedling stages are related to the trade-offs for trees ≥1 cm dbh. Here, we used principal components and correlation analyses to (1) determine the main demographic trade-offs among seed-to-seedling transition rates and growth and survival rates from the seedling to overstory size classes of 1188 tree species from large-scale forest dynamics plots in Panama, Puerto Rico, Ecuador, Taiwan, and Malaysia and (2) quantify the predictive power of maximum dbh, wood density, seed mass, and specific leaf area for species' position along these demographic trade-off gradients. In four out of five forests, the growth-survival trade-off was the most important demographic trade-off and encompassed growth and survival of both seedlings and trees ≥1 cm dbh. The second most important trade-off separated species with relatively fast growth and high survival at the seedling stage from species with relatively fast growth and high survival ≥1 cm dbh. The relationship between seed-to-seedling transition rates and these two trade-off aces differed between sites. All four traits were significant predictors for species' position along the two trade-off gradients, albeit with varying importance. We concluded that, after accounting for the species' position along the growth-survival trade-off, tree species tend to trade off growth and survival at the seedling with later life stages. This ontogenetic trade-off offers a mechanistic explanation for the stature-recruitment trade-off that constitutes an additional ontogenetic dimension of life-history variation in species-rich ecosystems., (© 2025 The Author(s). Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Forests sequester a substantial portion of anthropogenic carbon emissions. Many open questions concern how. We address two of these questions. Has leaf and fine litter production changed? And what is the contribution of old-growth forests? We address these questions with long-term records (≥10 years) of total, reproductive, and especially foliar fine litter production from 32 old-growth forests. We expect increases in forest productivity associated with rising atmospheric carbon dioxide concentrations and, in cold climates, with rising temperatures. We evaluate the statistical power of our analysis using simulations of known temporal trends parameterized with sample sizes (in number of years) and levels of interannual variation observed for each record. Statistical power is inadequate to detect biologically plausible trends for records lasting less than 20 years. Modest interannual variation characterizes fine litter production, and more variable phenomena will require even longer records to evaluate global change responses with sufficient statistical power. Just four old-growth forests have records of fine litter production lasting longer than 20 years, and these four provide no evidence for increases. Three of the four forests are in central Panama, also have long-term records of wood production, and both components of aboveground production are unchanged over 21-38 years. The possibility that recent increases in forest productivity are limited for old-growth forests deserves more attention., (Published 2025. This article is a U.S. Government work and is in the public domain in the USA.)

Premise: Soil microbes can influence patterns of diversity in plant communities via plant-soil feedbacks. Intraspecific plant-soil feedbacks occur when plant genotype leads to variations in soil microbial composition, resulting in differences in the performance of seedlings growing near their maternal plants versus seedlings growing near nonmaternal conspecific plants. How consistently such intraspecific plant-soil feedbacks occur in natural plant communities is unclear, especially in variable field conditions., Methods: In an in situ experiment with four native tree species on Barro Colorado Island (BCI), Panama, seedlings of each species were transplanted beneath their maternal tree or another conspecific tree in the BCI forest. Mortality and growth were assessed at the end of the wet season (~4 months post-transplant) and at the end of the experiment (~7 months post-transplant)., Results: Differences in seedling performance among field treatments were inconsistent among species and eroded over time. Effects of field environment were detected at the end of the wet season in two of the four species: Virola surinamensis seedlings had higher survival beneath their maternal tree than other conspecific trees, while seedling survival of Ormosia macrocalyx was higher under other conspecific trees. However, these differences were gone by the end of the experiment., Conclusions: Our results suggest that intraspecific plant-soil feedbacks may not be consistent in the field for tropical tree species and may have a limited role in determining seedling performance in tropical tree communities. Future studies are needed to elucidate the environmental and genetic factors that determine the incidence and direction of intraspecific plant-soil feedbacks in plant communities., (© 2024 The Authors. American Journal of Botany published by Wiley Periodicals LLC on behalf of Botanical Society of America.)

Future climate presents conflicting implications for forest biomass. We evaluate how plant hydraulic traits, elevated CO 2 levels, warming, and changes in precipitation affect forest primary productivity, evapotranspiration, and the risk of hydraulic failure. We used a dynamic vegetation model with plant hydrodynamics (FATES-HYDRO) to simulate the stand-level responses to future climate changes in a wet tropical forest in Barro Colorado Island, Panama. We calibrated the model by selecting plant trait assemblages that performed well against observations. These assemblages were run with temperature and precipitation changes for two greenhouse gas emission scenarios (2086-2100: SSP2-45, SSP5-85) and two CO 2 levels (contemporary, anticipated). The risk of hydraulic failure is projected to increase from a contemporary rate of 5.7% to 10.1-11.3% under future climate scenarios, and, crucially, elevated CO 2 provided only slight amelioration. By contrast, elevated CO 2 mitigated GPP reductions. We attribute a greater variation in hydraulic failure risk to trait assemblages than to either CO 2 or climate. Our results project forests with both faster growth (through productivity increases) and higher mortality rates (through increasing rates of hydraulic failure) in the neo-tropics accompanied by certain trait plant assemblages becoming nonviable., (© 2024 The Author(s). New Phytologist © 2024 New Phytologist Foundation.)

In tropical forests, trees strategically balance growth patterns to optimise fitness amid multiple environmental stressors. Wind poses the primary risk to a tree's mechanical stability, prompting developments such as thicker trunks to withstand the bending forces. Therefore, a trade-off in resource allocation exists between diameter growth and vertical growth to compete for light. We explore this trade-off by measuring the relative wind mortality risk for 95 trees in a tropical forest in Panama and testing how it varies with tree size, species and wind exposure. Surprisingly, local wind exposure and tree size had minimal impact on wind mortality risk; instead, species wood density emerged as the crucial factor. Low wood density species exhibited a significantly greater wind mortality risk, suggesting a prioritisation of competition for light over biomechanical stability. Our study highlights the pivotal role of wind safety in shaping the life-history strategy of trees and structuring diverse tropical forests., (© 2024 The Author(s). Ecology Letters published by John Wiley & Sons Ltd.)

Extending and safeguarding tropical forest ecosystems is critical for combating climate change and biodiversity loss. One of its constituents, lianas, is spreading and increasing in abundance on a global scale. This is particularly concerning as lianas negatively impact forests' carbon fluxes, dynamics, and overall resilience, potentially exacerbating both crises. While possibly linked to climate-change-induced atmospheric CO 2 elevation and drought intensification, the reasons behind their increasing abundance remain elusive. Prior research shows distinct physiological differences between lianas and trees, but it is unclear whether these differences confer a demographic advantage to lianas with climate change. Guided by extensive datasets collected in Panamanian tropical forests, we developed a tractable model integrating physiology, demography, and epidemiology. Our findings suggest that CO 2 fertilization, a climate change factor promoting forest productivity, gives lianas a demographic advantage. Conversely, factors such as extreme drought generally cause a decrease in liana prevalence. Such a decline in liana prevalence is expected from a physiological point of view because lianas have drought-sensitive traits. However, our analysis underscores the importance of not exclusively relying on physiological processes, as interactions with demographic mechanisms (i.e., the forest structure) can contrast these expectations, causing an increase in lianas with drought. Similarly, our results emphasize that identical physiological responses between lianas and trees still lead to liana increase. Even if lianas exhibit collinear but weaker responses in their performance compared to trees, a temporary liana prevalence increase might manifest driven by the faster response time of lianas imposed by their distinct life-history strategies than trees., Competing Interests: Competing interests statement:The authors declare no competing interest.

AbstractExplaining diversity in tropical forests remains a challenge in community ecology. Theory tells us that species differences can stabilize communities by reducing competition, while species similarities can promote diversity by reducing fitness differences and thus prolonging the time to competitive exclusion. Combined, these processes may lead to clustering of species such that species are niche differentiated across clusters and share a niche within each cluster. Here, we characterize this partial niche differentiation in a tropical forest in Panama by measuring spatial clustering of woody plants and relating these clusters to local soil conditions. We find that species were spatially clustered and the clusters were associated with specific concentrations of soil nutrients, reflecting the existence of nutrient niches. Species were almost twice as likely to recruit in their own nutrient niche. A decision tree algorithm showed that local soil conditions correctly predicted the niche of the trees with up to 85% accuracy. Iron, zinc, phosphorus, manganese, and soil pH were among the best predictors of species clusters.

Determining population demographic rates is fundamental to understanding differences in species' life-history strategies and their capacity to coexist. Calculating demographic rates, however, is challenging and requires long-term, large-scale censuses. Body size may serve as a simple predictor of demographic rate; can it act as a proxy for demographic rate when those data are unavailable? We tested the hypothesis that maximum body size predicts species' demographic rate using repeated censuses of the 77 most common liana species on the Barro Colorado Island, Panama (BCI) 50-ha plot. We found that maximum stem diameter does predict species' population turnover and demography. We also found that lianas on BCI can grow to the enormous diameter of 635 mm, indicating that they can store large amounts of carbon and compete intensely with tropical canopy trees. This study is the first to show that maximum stem diameter can predict plant species' demographic rates and that lianas can attain extremely large diameters. Understanding liana demography is particularly timely because lianas are increasing rapidly in many tropical forests, yet their species-level population dynamics remain chronically understudied. Determining per-species maximum liana diameters in additional forests will enable systematic comparative analyses of liana demography and potential influence across forest types., (© 2023 The Ecological Society of America.)

Premise: The specialized metabolites of plants are recognized as key chemical traits in mediating the ecology and evolution of sundry plant-biotic interactions, from pollination to seed predation. Intra- and interspecific patterns of specialized metabolite diversity have been studied extensively in leaves, but the diverse biotic interactions that contribute to specialized metabolite diversity encompass all plant organs. Focusing on two species of Psychotria shrubs, we investigated and compared patterns of specialized metabolite diversity in leaves and fruit with respect to each organ's diversity of biotic interactions., Methods: To evaluate associations between biotic interaction diversity and specialized metabolite diversity, we combined UPLC-MS metabolomic analysis of foliar and fruit specialized metabolites with existing surveys of leaf- and fruit-centered biotic interactions. We compared patterns of specialized metabolite richness and variance among vegetative and reproductive tissues, among plants, and between species., Results: In our study system, leaves interact with a far larger number of consumer species than do fruit, while fruit-centric interactions are more ecologically diverse in that they involve antagonistic and mutualistic consumers. This aspect of fruit-centric interactions was reflected in specialized metabolite richness-leaves contained more than fruit, while each organ contained over 200 organ-specific specialized metabolites. Within each species, leaf- and fruit-specialized metabolite composition varied independently of one another across individual plants. Contrasts in specialized metabolite composition were stronger between organs than between species., Conclusions: As ecologically disparate plant organs with organ-specific specialized metabolite traits, leaves and fruit can each contribute to the tremendous overall diversity of plant specialized metabolites., (© 2023 Botanical Society of America.)

Whether prey species avoid predators and predator species track prey is a poorly understood aspect of predator-prey interactions, given measuring prey tracking by predators and predator avoidance by prey is challenging. A common approach to study these interactions among mammals in field situations is to monitor the spatial proximity of animals at fixed times, using GPS tags fitted to individuals. However, this method is invasive and only allows tracking of a subset of individuals. Here, we use an alternative, noninvasive camera-trapping approach to monitor temporal proximity of predator and prey animals. We deployed camera traps at fixed locations on Barro Colorado Island, Panama, where the ocelot ( Leopardus pardalis ) is the principal mammalian predator, and tested two hypotheses: (1) prey animals avoid ocelots; and (2) ocelots track prey. We quantified temporal proximity of predators and prey by fitting parametric survival models to the time intervals between subsequent prey and predator captures by camera traps, and then compared the observed intervals to random permutations that retained the spatiotemporal distribution of animal activity. We found that time until a prey animal appeared at a location was significantly longer than expected by chance if an ocelot had passed, and that the time until an ocelot appeared at a location was significantly shorter than expected by chance after prey passage. These findings are indirect evidence for both predator avoidance and prey tracking in this system. Our results show that predator avoidance and prey tracking influence predator and prey distribution over time in a field setting. Moreover, this study demonstrates that camera trapping is a viable and noninvasive alternative to GPS tracking for studying certain predator-prey interactions., Competing Interests: The authors declare that they have no conflict of interest., (© The Author(s) 2022. Published by Oxford University Press on behalf of the American Society of Mammalogists.)

As currently circumscribed, Acrogenospora (Acrogenosporaceae, Minutisphaerales, Dothideomycetes) is a genus of saprobic hyphomycetes with distinctive conidia. Although considered common and cosmopolitan, the genus is poorly represented by sequence data, and no neotropical representatives are present in public sequence databases. Consequently, Acrogenospora has been largely invisible to ecological studies that rely on sequence-based identification. As part of an effort to identify fungi collected during ecological studies, we identified strains of Acrogenospora isolated in culture from seeds in the soil seed bank of a lowland tropical forest in Panama. Here we describe Acrogenospora terricola sp. nov. based on morphological and phylogenetic analyses. We confirm that the genus has a pantropical distribution. The observation of Acrogenospora infecting seeds in a terrestrial environment contrasts with previously described species in the genus, most of which occur on decaying wood in freshwater environments. This work highlights the often hidden taxonomic value of collections derived from ecological studies of fungal communities and the ways in which rich sequence databases can shed light on the identity, distributions and diversity of cryptic microfungi.

Identifying key traits that can serve as proxies for species drought resistance is crucial for predicting and mitigating the effects of climate change in diverse plant communities. Turgor loss point (π tlp ) is a recently emerged trait that has been linked to species distributions across gradients of water availability. However, a direct relationship between π tlp and species ability to survive drought has yet to be established for woody species. Using a manipulative field experiment to quantify species drought resistance (i.e., their survival response to drought), combined with measurements of π tlp for 16 tree species, we show a negative relationship between π tlp and seedling drought resistance. Using long-term forest plot data, we also show that π tlp predicts seedling survival responses to a severe El Niño-related drought, although additional factors are clearly also important. Our study demonstrates that species with lower π tlp exhibit higher survival under both experimental and natural drought. These results provide a missing cornerstone in the assessment of the traits underlying drought resistance in woody species and strengthen π tlp as a proxy for evaluating which species will lose or win under projections of exacerbating drought regimes., (© 2022 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Many well-studied animal species use conspicuous, repetitive signals that attract both mates and predators. Orthopterans (crickets, katydids, and grasshoppers) are renowned for their acoustic signals. In Neotropical forests, however, many katydid species produce extremely short signals, totaling only a few seconds of sound per night, likely in response to predation by acoustically orienting predators. The rare signals of these katydid species raises the question of how they find conspecific mates in a structurally complex rainforest. While acoustic mechanisms, such as duetting, likely facilitate mate finding, we test the hypothesis that mate finding is further facilitated by colocalization on particular host plant species. DNA barcoding allows us to identify recently consumed plants from katydid stomach contents. We use DNA barcoding to test the prediction that katydids of the same species will have closely related plant species in their stomach. We do not find evidence for dietary specialization. Instead, katydids consumed a wide mix of plants within and across the flowering plants (27 species in 22 genera, 16 families, and 12 orders) with particular representation in the orders Fabales and Laurales. Some evidence indicates that katydids may gather on plants during a narrow window of rapid leaf out, but additional investigations are required to determine whether katydid mate finding is facilitated by gathering at transient food resources., Competing Interests: Conflicts of Interest: The authors declare no conflict of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript, or in the decision to publish the results.

For community ecologists, "neutral or not?" is a fundamental question, and thus, rejecting neutrality is an important first step before investigating the deterministic processes underlying community dynamics. Hubbell's neutral model is an important contribution to the exploration of community dynamics, both technically and philosophically. However, the neutrality tests for this model are limited by a lack of statistical power, partly because the zero-sum assumption of the model is unrealistic. In this study, we developed a neutrality test for local communities that implements non-zero-sum community dynamics and determines the number of new species ( N sp ) between observations. For the non-zero-sum neutrality test, the model distributed the expected N sp , as calculated by extensive simulations, which allowed us to investigate the neutrality of the observed community by comparing the observed N sp with distributions of the expected N sp derived from the simulations. For this comparison, we developed a new "non-zero-sum N sp test," which we validated by running multiple neutral simulations using different parameter settings. We found that the non-zero-sum N sp test rejected neutrality at a near-significance level, which justified the validity of our approach. For an empirical test, the non-zero-sum N sp test was applied to real tropical tree communities in Panama and Malaysia. The non-zero-sum N sp test rejected neutrality in both communities when the observation interval was long and N sp was large. Hence, the non-zero-sum N sp test is an effective way to examine neutrality and has reasonable statistical power to reject the neutral model, especially when the observed N sp is large. This unique and simple approach is statistically powerful, even though it only employs two temporal sequences of community data. Thus, this test can be easily applied to existing datasets. In addition, application of the test will provide significant benefits for detecting changing biodiversity under climate change and anthropogenic disturbance., Competing Interests: The authors declare no conflict of interest., (© 2022 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Bees, ants, and wasps are well known to visually navigate when traveling between their nests and foraging sites. When leaving their nest, landmarks in the vicinity are memorized and used upon return to locate the nest entrance. 1 , 2 The Neotropical nocturnal sweat bee Megalopta genalis navigates under the forest canopy at light intensities ten times dimmer than starlight. 3 Despite these dim conditions, Megalopta is able to memorize visual landmarks around the nest entrance in the frontal visual field. 4 Even though frontal landmarks can clearly be discerned by Megalopta, the visual feature of greatest contrast in the rainforest at night is actually the dark dorsal silhouette of the distant canopy against the brighter night sky. Several species of ants, 5-10 as well as a subsocial shield bug, 11 use bright open gaps in the canopy as dorsal landmarks to navigate home while walking. Here we show that Megalopta is also able to distinguish dorsal landmarks during homing, the first flying insect known with this capacity. Megalopta is able to discriminate between differently oriented dorsal black striped patterns, or an "artificial canopy" of black circles, and to use this information to locate its nest entrance. These results suggest that the local foliage patterns created by the canopy against the brighter sky could potentially provide the bee with reliable landmark information for navigation during foraging and homing at night. VIDEO ABSTRACT., Competing Interests: Declaration of interests The authors declare that they have no competing interests., (Copyright © 2021 The Author(s). Published by Elsevier Inc. All rights reserved.)

Seeds collected from trees, shrubs and lianas growing on Barro Colorado Island, Panama, were analyzed for their content of phenolic compounds, oxidative activities and protein precipitation capacities. Proanthocyanidins and hydrolysable tannins were detected in one-third of 189 studied species. The most oxidatively active group of species were the ones containing prodelphinidins and ellagitannins whereas the species that had the highest protein precipitation capacity in relation to their total phenolics were the ones containing punicalagin. In addition, the oxidative activity and relative protein precipitation capacity were exceptionally high in the proanthocyanidin-rich genus Psychotria. This study offers a comprehensive overview on the tannin composition and the alkaline oxidative activities and protein precipitation capacities of the seeds of tropical plants., (Copyright © 2021 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Predation produces intense selection and a diversity of defences. Reactive defences are triggered by predator cues, whereas proactive defences are always in effect. We assess whether prey rely on proactive defences when predator cues do not correlate well with predation risk. Many bats use echolocation to hunt insects, and many insects have evolved to hear bats. However, in species-rich environments like Neotropical forests, bats have extremely diverse foraging strategies, and the presence of echolocation corresponds only weakly to the presence of predators. We assess whether katydids that live in habitats with many non-dangerous bat species stop calling when exposed to echolocation. For 11 species of katydids, we quantified behavioural and neural responses to predator cues, and katydid signalling activity over 24 h periods. Despite having the sensory capacity to detect predators, many Neotropical forest katydids continued calling in the presence of predator cues, displaying proactive defences instead (short, infrequent calls totalling less than 2 cumulative seconds of sound per 24 h). Neotropical katydid signalling illustrates a fascinating case where trophic interactions are probably mediated by a third group: bats with alternative foraging strategies (e.g. frugivory). Although these co-occurring bats are not trophically connected, their mere presence disrupts the correlation between cue and predation risk.

Tropical forests are a key determinant of the functioning of the Earth system, but remain a major source of uncertainty in carbon cycle models and climate change projections. In this study, we present an updated land model (LM3PPA-TV) to improve the representation of tropical forest structure and dynamics in Earth system models (ESMs). The development and parameterization of LM3PPA-TV drew on extensive datasets on tropical tree traits and long-term field censuses from Barro Colorado Island (BCI), Panama. The model defines a new plant functional type (PFT) based on the characteristics of shade-tolerant, tropical tree species, implements a new growth allocation scheme based on realistic tree allometries, incorporates hydraulic constraints on biomass accumulation, and features a new compartment for tree branches and branch fall dynamics. Simulation experiments reproduced observed diurnal and seasonal patterns in stand-level carbon and water fluxes, as well as mean canopy and understory tree growth rates, tree size distributions, and stand-level biomass on BCI. Simulations at multiple sites captured considerable variation in biomass and size structure across the tropical forest biome, including observed responses to precipitation and temperature. Model experiments suggested a major role of water limitation in controlling geographic variation forest biomass and structure. However, the failure to simulate tropical forests under extreme conditions and the systematic underestimation of forest biomass in Paleotropical locations highlighted the need to incorporate variation in hydraulic traits and multiple PFTs that capture the distinct floristic composition across tropical domains. The continued pressure on tropical forests from global change demands models which are able to simulate alternative successional pathways and their pace to recovery. LM3PPA-TV provides a tool to investigate geographic variation in tropical forests and a benchmark to continue improving the representation of tropical forests dynamics and their carbon storage potential in ESMs., (© 2020 John Wiley & Sons Ltd.)

Tropical forests challenge us to understand biodiversity, as numerous seemingly similar species persist on only a handful of shared resources. Recent ecological theory posits that biodiversity is sustained by a combination of species differences reducing interspecific competition and species similarities increasing time to competitive exclusion. Together, these mechanisms counterintuitively predict that competing species should cluster by traits, in contrast with traditional expectations of trait overdispersion. Here, we show for the first time that trees in a tropical forest exhibit a clustering pattern. In a 50-ha plot on Barro Colorado Island in Panama, species abundances exhibit clusters in two traits connected to light capture strategy, suggesting that competition for light structures community composition. Notably, we find four clusters by maximum height, quantitatively supporting the classical grouping of Neotropical woody plants into shrubs, understory, midstory, and canopy layers., (© 2020 by the Ecological Society of America.)

Plant enemies that attack chemically similar host species are thought to mediate competitive exclusion of chemically similar plants and select for chemical divergence among closely related species. This hypothesis predicts that plant defenses should diverge rapidly, minimizing phylogenetic signal. To evaluate this prediction, we quantified metabolomic similarity for 203 tree species that represent >89% of all individuals in large forest plots in Maryland and Panama. We constructed molecular networks based on mass spectrometry of all 203 species, quantified metabolomic similarity for all pairwise combinations of species, and used phylogenetically independent contrasts to evaluate how pairwise metabolomic similarity varies phylogenetically. Leaf metabolomes exhibited clear phylogenetic signal for the temperate plot, which is inconsistent with the prediction. In contrast, leaf metabolomes lacked phylogenetic signal for the tropical plot, with particularly low metabolomic similarity among congeners. In addition, community-wide variation in metabolomes was much greater for the tropical community, with single tropical genera supporting greater metabolomic variation than the entire temperate community. Our results are consistent with the hypothesis that stronger plant-enemy interactions lead to more rapid divergence and greater metabolomic variation in tropical than temperate plants. Additional community-level foliar metabolomes will be required from tropical and temperate forests to evaluate this hypothesis., (© 2018 by the Ecological Society of America.)

Conspecific negative density dependence (CNDD) is thought to promote plant species diversity. Theoretical studies showing the importance of CNDD often assumed that all species are equally susceptible to CNDD; however, recent empirical studies have shown species can differ greatly in their susceptibility to CNDD. Using a theoretical model, we show that interspecific variation in CNDD can dramatically alter its impact on diversity. First, if the most common species are the least regulated by CNDD, then the stabilising benefit of CNDD is reduced. Second, when seed dispersal is limited, seedlings that are susceptible to CNDD are at a competitive disadvantage. When parameterised with estimates of CNDD from a tropical tree community in Panama, our model suggests that the competitive inequalities caused by interspecific variation in CNDD may undermine many species' ability to persist. Thus, our model suggests that variable CNDD may make communities less stable, rather than more stable., (© 2018 John Wiley & Sons Ltd/CNRS.)

Seeds of tropical pioneer trees have chemical and physical characteristics that determine their capacity to persist in the soil seed bank. These traits allow seeds to survive in the soil despite diverse predators and pathogens, and to germinate and recruit even decades after dispersal. Defenses in seedlings and adult plants often are described in terms of tradeoffs between chemical and physical defense, but the interplay of defensive strategies has been evaluated only rarely for seeds. Here we evaluated whether classes of seed defenses were negatively correlated across species (consistent with tradeoffs in defense strategies), or whether groups of traits formed associations across species (consistent with seed defense syndromes). Using 16 of the most common pioneer tree species in a neotropical lowland forest in Panama we investigated relationships among four physical traits (seed fracture resistance, seed coat thickness, seed permeability, and seed mass) and two chemical traits (number of phenolic compounds and phenolic peak area), and their association with seed persistence. In addition, seed toxicity was assessed with bioassays in which we evaluated the activity of seed extracts against representative fungal pathogens and a model invertebrate. We did not find univariate tradeoffs between chemical and physical defenses. Instead, we found that seed permeability - a trait that distinguishes physical dormancy from other dormancy types - was positively associated with chemical defense traits and negatively associated with physical defense traits. Using a linear discriminant analysis and a hierarchical cluster analysis we found evidence to distinguish three distinct seed defense syndromes that correspond directly with seed dormancy classes (i.e., quiescent, physical, and physiological). Our data suggest that short and long-term persistence of seeds can be achieved via two strategies: having permeable seeds that are well defended chemically, corresponding to the physiologically dormant defense syndrome; or having impermeable seeds that are well defended physically, corresponding to the physically dormant defense syndrome. In turn, transient seeds appear to have a lower degree of chemical and physical defenses, corresponding to the quiescent defense syndrome. Overall, we find that seed defense and seed dormancy are linked, suggesting that environmental pressures on seed persistence and for delayed germination can select for trait combinations defining distinct dormancy-defense syndromes., (© 2018 by the Ecological Society of America.)

Life-history theory posits that trade-offs between demographic rates constrain the range of viable life-history strategies. For coexisting tropical tree species, the best established demographic trade-off is the growth-survival trade-off. However, we know surprisingly little about co-variation of growth and survival with measures of reproduction. We analysed demographic rates from seed to adult of 282 co-occurring tropical tree and shrub species, including measures of reproduction and accounting for ontogeny. Besides the well-established fast-slow continuum, we identified a second major dimension of demographic variation: a trade-off between recruitment and seedling performance vs. growth and survival of larger individuals (≥ 1 cm dbh) corresponding to a 'stature-recruitment' axis. The two demographic dimensions were almost perfectly aligned with two independent trait dimensions (shade tolerance and size). Our results complement recent analyses of plant life-history variation at the global scale and reveal that demographic trade-offs along multiple axes act to structure local communities., (© 2018 The Authors. Ecology Letters published by CNRS and John Wiley & Sons Ltd.)

Metabarcoding of vertebrate DNA derived from carrion flies has been proposed as a promising tool for biodiversity monitoring. To evaluate its efficacy, we conducted metabarcoding surveys of carrion flies on Barro Colorado Island (BCI), Panama, which has a well-known mammal community, and compared our results against diurnal transect counts and camera trapping. We collected 1,084 flies in 29 sampling days, conducted metabarcoding with mammal-specific (16S) and vertebrate-specific (12S) primers, and sequenced amplicons on Illumina MiSeq. For taxonomic assignment, we compared blast with the new program protax, and we found that protax improved species identifications. We detected 20 mammal, four bird, and one lizard species from carrion fly metabarcoding, all but one of which are known from BCI. Fly metabarcoding detected more mammal species than concurrent transect counts (29 sampling days, 13 species) and concurrent camera trapping (84 sampling days, 17 species), and detected 67% of the number of mammal species documented by 8 years of transect counts and camera trapping combined, although fly metabarcoding missed several abundant species. This study demonstrates that carrion fly metabarcoding is a powerful tool for mammal biodiversity surveys and has the potential to detect a broader range of species than more commonly used methods., (© 2017 John Wiley & Sons Ltd.)

Multiple anthropogenic drivers affect every natural community, and there is broad interest in using functional traits to understand and predict the consequences for future biodiversity. There is, however, no consensus regarding the choice of analytical methods. We contrast species- and community-level analyses of change in the functional composition for four traits related to drought tolerance using three decades of repeat censuses of trees in the 50-ha Forest Dynamics Plot on Barro Colorado Island, Panama. Community trait distributions shifted significantly through time, which may indicate a shift toward more drought tolerant species. However, at the species level, changes in abundance were unrelated to trait values. To reconcile these seemingly contrasting results, we evaluated species-specific contributions to the directional shifts observed at the community level. Abundance changes of just one to six of 312 species were responsible for the community-level shifts observed for each trait. Our results demonstrate that directional changes in community-level functional composition can result from idiosyncratic change in a few species rather than widespread community-wide changes associated with functional traits. Future analyses of directional change in natural communities should combine community-, species-, and possibly individual-level analyses to uncover relationships with function that can improve understanding and enable prediction., (© 2017 by the Ecological Society of America.)

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.)

Contagious seed dispersal refers to the tendency for some sites to receive many dispersed seeds while other sites receive few dispersed seeds. Contagious dispersal can lead to interspecific associations in seed arrival, and this in turn might lead to interspecific associations in seedling recruitment. We evaluate the extent of spatially contagious seed arrival, the frequency of positive interspecific associations in seed arrival, and their consequences for seedling recruitment at the community level in a tropical moist forest. We quantified seed arrival to 200 passive seed traps for 28 yr of weekly censuses and seedling recruitment to 600 1-m 2 quadrats for 21 yr of annual censuses on Barro Colorado Island, Panama. We assessed whether spatially contagious seed dispersal was more important among zoochorous species than among anemochorous species, increased in importance with similarity in fruiting times, and led to interspecific associations in seed arrival and seedling recruitment. We controlled adult seed source associations statistically to evaluate predicted relationships. We found that spatially contagious seed arrival was widespread among zoochorous species, but also occurred among anemochorous species when the strong, consistent trade winds were present. Significant interspecific associations in seed arrival were more likely for pairs of species with zoochorous seeds and similar fruiting times and persisted through seedling recruitment. Thus, interspecifically contagious seed dispersal affects local species composition and alters the mixture of interspecific interactions through the seed, germination, and early seedling stages in this forest. Future investigations should consider the implications of interspecific association at the regeneration stages documented here for later life stages and species coexistence., (© 2016 by the Ecological Society of America.)

Herbaceous plants are a key component of tropical forests. Previous work indicates that herbs contribute substantially to the species richness of tropical plant communities. However, the processes structuring tropical herb diversity, and how they contrast with woody communities, have been underexplored. Within the understory of a 50-ha forest dynamics plot in central Panama, we compared the diversity, distribution, and abundance of vascular herbaceous plants with woody seedlings (i.e., tree and lianas <1 cm DBH and ≥20 cm tall). Beta-diversity was calculated for each community using a null model approach. We then assessed the similarity in alpha and beta-diversity among herbs, tree seedlings, and liana seedlings. Strengths of habitat associations were measured using permutational ANOVA among topographic habitat-types. Variance partitioning was then used to quantify the amount of variation in species richness and composition explained by spatial and environmental variables (i.e., topography, soils, and shade) for each growth form. Species richness and diversity were highest for tree seedlings, followed by liana seedlings and then herbs. In contrast, beta-diversity was 16-127% higher for herbs compared to woody seedlings, indicating higher spatial variation in this stratum. We observed no correlation between local richness or compositional uniqueness of herbs and woody seedlings across sites, indicating that different processes control the spatial patterns of woody and herbaceous diversity and composition. Habitat associations were strongest for herbs, as indicated by greater compositional dissimilarity among habitat types. Likewise, environmental variables explained a larger proportion of the variation in species richness and composition for herbs than for woody seedlings (richness = 25%, 14%, 12%; composition = 25%, 9%, 6%, for herbs, trees, and lianas, respectively). These differences between strata did not appear to be due to differences in lifespan alone, based on data from adult trees. Our results point to contrasting assembly mechanisms for herbaceous and woody communities, with herbs showing stronger niche-derived structure. Future research on tropical herbaceous communities is likely to yield new insights into the many processes structuring diverse plant communities., (© 2016 by the Ecological Society of America.)

Background: Nematodes are a very diverse and extremely abundant group of animals, but their occurrence in the tropics is surprisingly little understood. We investigated the meiofauna of epiphytic tank bromeliads in the lowlands of Panama with particular emphasis on nematodes., Results: We encountered 89 morphospecies of nematodes in 54 bromeliad tanks, which were sampled in the wet and the dry season. Rotifers were by far the most abundant group in both the dry and the wet season (with up to 960 individual ml(-1)), followed by nematodes, annelids and harpacticoid copepods. Individual plants hosted up to 25 nematode species. These nematodes represented a diversity of feeding guilds, suction-feeders and deposit-feeders being most abundant. The relative abundances of feeding-types of nematodes differed considerably in the wet and dry season. Both species richness and abundance were strongly correlated with the size of the phytotelms and the season, while species diversity assessed with the Shannon-index was affected by neither of the two., Conclusion: This is the first study with a particular focus on the diversity of nematodes in tank bromeliads. We document a meiofauna of considerable abundance and diversity, which suggests important functional roles in ecological processes such as decomposition, which in turn warrants further study.

Germination from the soil seed bank (SSB) is an important determinant of species composition in tropical forest gaps, with seed persistence in the SSB allowing trees to recruit even decades after dispersal. The capacity to form a persistent SSB is often associated with physical dormancy, where seed coats are impermeable at the time of dispersal. Germination literature often speculates, without empirical evidence, that dormancy-break in physically dormant seeds is the result of microbial action and/or abrasion by soil particles. We tested the microbial/soil abrasion hypothesis in four widely distributed neotropical pioneer tree species (Apeiba membranacea, Luehea seemannii, Ochroma pyramidale, and Cochlospermum vitifolium). Seeds were buried in five common gardens in a lowland tropical forest in Panama, and recovered at 1, 3, 6, and 12 months after burial. Seed permeability, microbial infection, seed coat thickness, and germination were measured. Parallel experiments compared the germination fraction of fresh and aged seeds without soil contact, and in seeds as a function of seed permeability. Contrary to the microbial/soil abrasion hypothesis the proportion of permeable seeds, and of seeds infected by cultivable microbes, decreased as a function of burial duration. Furthermore, seeds stored in dark and dry conditions for 2 years showed a higher proportion of seed germination than fresh seeds in identical germination conditions. We determined that permeable seeds of A. membranacea and O. pyramidale had cracks in the chalazal area or lacked the chalazal plug, whereas all surfaces of impermeable seeds were intact. Our results are inconsistent with the microbial/soil abrasion hypothesis of dormancy loss and instead suggest the existence of multiple dormancy phenotypes, where a fraction of each seed cohort is dispersed in a permeable state and germinates immediately, while the impermeable seed fraction accounts for the persistent SSB. Thus, we conclude that fluctuations in the soil temperature in the absence of soil abrasion and microbial infection are sufficient to break physical dormancy on seeds of tropical pioneer trees.

Treefall gaps play an important role in tropical forest dynamics and in determining above-ground biomass (AGB). However, our understanding of gap disturbance regimes is largely based either on surveys of forest plots that are small relative to spatial variation in gap disturbance, or on satellite imagery, which cannot accurately detect small gaps. We used high-resolution light detection and ranging data from a 1500 ha forest in Panama to: (i) determine how gap disturbance parameters are influenced by study area size, and the criteria used to define gaps; and (ii) to evaluate how accurately previous ground-based canopy height sampling can determine the size and location of gaps. We found that plot-scale disturbance parameters frequently differed significantly from those measured at the landscape-level, and that canopy height thresholds used to define gaps strongly influenced the gap-size distribution, an important metric influencing AGB. Furthermore, simulated ground surveys of canopy height frequently misrepresented the true location of gaps, which may affect conclusions about how relatively small canopy gaps affect successional processes and contribute to the maintenance of diversity. Across site comparisons need to consider how gap definition, scale and spatial resolution affect characterizations of gap disturbance, and its inferred importance for carbon storage and community composition.

Although competition between social groups is central to hypotheses about the evolution of human social organization, competitive interactions among group-mates are thought to play a more dominant role in shaping the behavior and ecology of other primate species. However, few studies have directly tested the impact of intergroup conflicts in non-human primates. What is the cost of defeat? To address this question, the movements of six neighboring white-faced capuchin (Cebus capucinus) social groups living on Barro Colorado Island, Panama were tracked simultaneously using an Automated Radio Telemetry System (ARTS), for a period of six months. Groups moved 13% (441 m) further on days they lost interactions compared with days they won interactions. To cover these larger distances, they traveled faster, stopped less frequently, and remained active later in the evening. Defeat also caused groups to alter their patterns of space use. Losing groups had straighter travel paths than winning groups, larger net displacements and were more likely to change their sleeping site. These results demonstrate that losing groups pay increased travel costs and suggest that they forage in low-quality areas. They provide some of the first direct evidence that intergroup conflicts have important energetic consequences for members of competitively unsuccessful primate social groups. A better understanding of how intergroup competition impacts patterns of individual fitness is thus needed to clarify the role that this group-level process plays in shaping the evolution of human- and non-human primate behavior., (Copyright © 2013 Wiley Periodicals, Inc.)

Fifty years ago, Baker and Fedorov proposed that the high species diversity of tropical forests could arise from the combined effects of inbreeding and genetic drift leading to population differentiation and eventually to sympatric speciation. Decades of research, however have failed to support the Baker-Fedorov hypothesis (BFH), and it has now been discarded in favor of a paradigm where most trees are self-incompatible or strongly outcrossing, and where long-distance pollen dispersal prevents population drift. Here, we propose that several hyper-diverse genera of tropical herbs and shrubs, including Piper (>1,000 species), may provide an exception. Species in this genus often have aggregated, high-density populations with self-compatible breeding systems; characteristics which the BFH would predict lead to high local genetic differentiation. We test this prediction for five Piper species on Barro Colorado Island, Panama, using Amplified Fragment Length Polymorphism (AFLP) markers. All species showed strong genetic structure at both fine- and large-spatial scales. Over short distances (200-750 m) populations showed significant genetic differentiation (Fst 0.11-0.46, P < 0.05), with values of spatial genetic structure that exceed those reported for other tropical tree species (Sp = 0.03-0.136). This genetic structure probably results from the combined effects of limited seed and pollen dispersal, clonal spread, and selfing. These processes are likely to have facilitated the diversification of populations in response to local natural selection or genetic drift and may explain the remarkable diversity of this rich genus.

Changes in abscisic acid (ABA) content and several water relations parameters were studied in the epiphytic orchid Dimerandra emarginata in the tropical lowlands of Barro Colorado Island, Panama. Similar to previous observations of size dependency of physiological parameters in vascular epiphytes, we found significant differences in ABA accumulation (both in terms of temporal patterns and amount) in small and large plants under drought stress. The highest values of ABA content in leaves, stems, and roots were found in the smaller specimens, reaching the levels of poikilohydric resurrection plants. Substantial accumulation of ABA occurred after stomatal closure, indicating other functions of ABA than stomatal regulation in this bark epiphyte.

We present an analysis of the long-term survival of two cohorts of seedlings of the tropical canopy tree Ocotea whitei (Lauraceae) on a 1-ha plot of mature, lowland moist forest on Barro Colorado Island, Panamá. In 1980, we counted an even-aged cohort of seedlings that germinated in 1979, then measured and tagged survivors in 1981. We also measured and tagged a second, smaller cohort of seedlings that germinated in 1981. We followed the subsequent survival of all seedlings through 1985. Seedling mortality was phenotypically, temporally, and spatially non-random. Important correlates of non-random mortality included: (1) seedling size and age, (2) an El Niño drought, and (3) biotic neighborhood. Larger and older seedlings survived better than smaller and younger seedlings, respectively, and the El Niño-related drought of 1982-1983 was associated with elevated mortality rates. Seedling density, which was strongly correlated with the proximity to the nearest conspecific adult, increased mortality. The observed mortality patterns suggest that processes consistent with the Janzen-Connell hypothesis operate during the recruitment phase of O. whitei population dynamics. However, the processes causing the observed density- and distance-dependent mortality may vary with factors such as total seed number, seedling size, and climatic variation, making it difficult to determine whether time-integrated seedling-to-adult spacing mechanisms other than self-thinning operate on a given plant population. After 6 years in the hectare studied, survivors remained densest and most numerous underneath the adult trees. We conclude that only long-term demographic data, collected at a variety of scales on a variety of species, will ultimately answer the question: do Janzen-Connell effects contribute substantially to structuring tropical forests?

Photosynthetic performance under dynamic light regimes was assessed in four different species of tropical shrubs from the family Rubiaceae via field gas exchange measurements conducted on Barro Colorado Island, Panamá. Rates of photosynthetic induction and induction loss were assessed throughout the day in both the wet and dry seasons in order to determine the relative roles of stomata and biochemistry in limiting photosynthetic performance under transient light conditions. A high degree of coordination was observed between stomatal conductance and biochemical capacity for CO 2 assimilation during induction. Rates of biochemical and overall photosynthetic induction sharply decreased when initial stomatal conductance fell below a narrow range of critical values. Time of day or season did not affect rates of biochemical deactivation upon shading, but did influence stomatal closure, which often exerted a significant influence over induction loss in the darkness. In measurements of total assimilation due to a 60-s light pulse, both biochemical activity and stomatal conductance were linearly related to total CO 2 uptake. Only during the mornings of the wet season was stomatal conductance consistently high enough to be non-limiting to dynamic photosynthetic performance. At all other times, stomatal behavior exercised significant influence over induction times, photosynthetic induction loss, and total CO 2 uptake from 60-s light pulses.

Rates of photosynthetic induction upon exposure to high light and rates of induction loss after darkening the leaf were measured in the field for four species of tropical shrubs in the family Rubiaceae. During wet season mornings, stomatal conductance (g s ) in the shade prior to induction was generally high enough so that the time course of induction was determined primarily by rates of activation of biochemical processes. During wet season afternoons, however, g s values in the shade tended to be considerably lower and photosynthetic induction following a light increase exhibited a slower time course. In the afternoon, the time course of induction was determined by a combination of stomatal opening time and the rates of activation of light regulated enzymes. Stomatal behavior was also correlated with patterns of induction loss following a transfer from high light to darkness. In the afternoon, maximum g s was lower for all species, and for a given time in the darkness, leaves showed a greater loss of induction in the afternoon than in the morning. During the dry season, maximum g s and average values for g s in the shade were reduced in all species. Along with these shifts in stomatal behavior, reduced rates of photosynthetic induction were observed. In the high-light species, the lower maximum g s values observed during the dry season were also correlated with increased induction loss for a given time in the darkness. For all species, stomatal behavior was affected by season and time of day and, with the exception of wet season mornings, stomata appeared to exert significant control over rates of induction and patterns of induction loss. The results of simulation modeling suggest that the observed seasonal and diurnal changes in rates of induction and induction loss can have significant consequences on sunfleck carbon gain under a dynamic light regime.

The occurrence of crassulacean acid metabolism (CAM) among the epiphyte flora of the lowland forest on Barro Colorado Island (BCJ), Panama, was investigated. A total of 116 species was included, i.e. about 2/3 of the known epiphyte taxa. As judged from the carbon isotope ratios and the absence of Kranz anatomy, indications of CAM were found in 29 species of three families, Orchidaceae (20), Bromeliaceae (7), and Cactaceae (2). We estimate that about 25% of the epiphyte flora of BCI are CAM plants. CAM was most prevalent in exposed sites, but even in the understorey two epiphyte species engage in CAM.

Pre-dawn xylem pressures were measured with bubble manometers attached near the stem bases of 32 species of vines on Barro Colorado Island, Panama, to determine if pressures were sufficient to allow for possible refilling of embolized vessels. Of 29 dicotyledonous species 26 exhibited only negative xylem pressures, even pre-dawn during the wet season. In contrast, three members of the Dilleniaceae exhibited positive pre-dawn xylem pressures, with a maximum of 64 kPa in Doliocarpusmajor. A pressure of 64 kPa is sufficient to push water to a height of 6.4 m against gravity, but the specimens reached heights of 18 m. Thus, in all 29 dicotyledons examined, the xylem pressures were not sufficient to refill embolized vessels in the upper stems. In contrast, two of the smaller, non-dicotyledonous vines, the climbing fern Lygodiumvenustrum and the viny bamboo Rhipidocladumracemiflorum, had xylem pressures sufficient to push water to the apex of the plants. Therefore, a root pressure mechanism to reverse embolisms in stem xylem could apply to some but not to most of the climbing plants that were studied.