Wind Shapes the Growth Strategies of Trees in a Tropical Forest.

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.

Soils and topography control natural disturbance rates and thereby forest structure in a lowland tropical landscape.

Tree mortality is a major control over tropical forest carbon stocks globally but the strength of associations between abiotic drivers and tree mortality within forested landscapes is poorly understood. Here, we used repeat drone photogrammetry across 1500 ha of forest in Central Panama over 5 years to quantify spatial variation in canopy disturbance rates and its predictors. We identified 11,153 canopy disturbances greater than 25 m2 in area, including treefalls, large branchfalls and standing dead trees, affecting 1.9% of area per year. Soil type, forest age and topography explained up to 46%-67% of disturbance rate variation at spatial grains of 58-64 ha, with higher rates in older forests, steeper slopes and local depressions. Furthermore, disturbance rates predicted the proportion of low canopy area across the landscape, and mean canopy height in old growth forests. Thus abiotic factors drive variation in disturbance rates and thereby forest structure at landscape scales.

Leaf turgor loss point shapes local and regional distributions of evergreen but not deciduous tropical trees.

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'; πtlp ), 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 πtlp were associated with drier habitats, with πtlp explaining 28% and 32% of habitat association on local and regional scales, respectively, greatly exceeding the predictive power of WD and LMA. In contrast, πtlp did not predict habitat associations among deciduous species. Across spatial scales, πtlp 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.

Patterns and mechanisms of spatial variation in tropical forest productivity, woody residence time, and biomass.

Tropical forests vary widely in biomass carbon (C) stocks and fluxes even after controlling for forest age. A mechanistic understanding of this variation is critical to accurately predicting responses to global change. We review empirical studies of spatial variation in tropical forest biomass, productivity and woody residence time, focusing on mature forests. Woody productivity and biomass decrease from wet to dry forests and with elevation. Within lowland forests, productivity and biomass increase with temperature in wet forests, but decrease with temperature where water becomes limiting. Woody productivity increases with soil fertility, whereas residence time decreases, and biomass responses are variable, consistent with an overall unimodal relationship. Areas with higher disturbance rates and intensities have lower woody residence time and biomass. These environmental gradients all involve both direct effects of changing environments on forest C fluxes and shifts in functional composition - including changing abundances of lianas - that substantially mitigate or exacerbate direct effects. Biogeographic realms differ significantly and importantly in productivity and biomass, even after controlling for climate and biogeochemistry, further demonstrating the importance of plant species composition. Capturing these patterns in global vegetation models requires better mechanistic representation of water and nutrient limitation, plant compositional shifts and tree mortality.

Integrating high resolution drone imagery and forest inventory to distinguish canopy and understory trees and quantify their contributions to forest structure and dynamics.

Tree growth and survival differ strongly between canopy trees (those directly exposed to overhead light), and understory trees. However, the structural complexity of many tropical forests makes it difficult to determine canopy positions. The integration of remote sensing and ground-based data enables this determination and measurements of how canopy and understory trees differ in structure and dynamics. Here we analyzed 2 cm resolution RGB imagery collected by a Remotely Piloted Aircraft System (RPAS), also known as drone, together with two decades of bi-annual tree censuses for 2 ha of old growth forest in the Central Amazon. We delineated all crowns visible in the imagery and linked each crown to a tagged stem through field work. Canopy trees constituted 40% of the 1244 inventoried trees with diameter at breast height (DBH) > 10 cm, and accounted for ~70% of aboveground carbon stocks and wood productivity. The probability of being in the canopy increased logistically with tree diameter, passing through 50% at 23.5 cm DBH. Diameter growth was on average twice as large in canopy trees as in understory trees. Growth rates were unrelated to diameter in canopy trees and positively related to diameter in understory trees, consistent with the idea that light availability increases with diameter in the understory but not the canopy. The whole stand size distribution was best fit by a Weibull distribution, whereas the separate size distributions of understory trees or canopy trees > 25 cm DBH were equally well fit by exponential and Weibull distributions, consistent with mechanistic forest models. The identification and field mapping of crowns seen in a high resolution orthomosaic revealed new patterns in the structure and dynamics of trees of canopy vs. understory at this site, demonstrating the value of traditional tree censuses with drone remote sensing.

Allometric constraints and competition enable the simulation of size structure and carbon fluxes in a dynamic vegetation model of tropical forests (LM3PPA-TV).

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.

No bejel among Surinamese, Antillean and Dutch syphilis diagnosed patients in Amsterdam between 2006-2018 evidenced by multi-locus sequence typing of Treponema pallidum isolates.

BACKGROUND: Treponema pallidum subspecies pallidum (TPA) and subsp. endemicum (TEN) are the causative agents of syphilis and bejel, respectively. TEN shows similar clinical manifestations and is morphologically and serologically indistinguishable from TPA. Recently, bejel was found outside of its assumed endemic areas. Using molecular typing we aimed to discover bejel and characterize circulating TPA types among syphilis cases with Surinamese, Antillean and Dutch ethnicity in Amsterdam. METHODS: DNA was extracted from 137 ulcer swabs, which tested positive in the in-house diagnostic PCR targeting the polA gene. Samples were collected between 2006 and 2018 from Surinamese, Antillean and Dutch patients attending the Amsterdam STI clinic. Multilocus sequence typing was performed by partial sequence analysis of the tp0136, tp0548 and tp0705 genes. In addition, the 23S rRNA loci were analyzed for A2058G and A2059G macrolide resistance mutations. RESULTS: We found 17 distinct allelic profiles in 103/137 (75%) fully typed samples, which were all TPA and none TEN. Of the strains, 82.5% were SS14-like and 17.5% Nichols-like. The prevalence of Nichols-like strains found in this study is relatively high compared to nearby countries. The most prevalent types were 1.3.1 (42%) and 1.1.1 (19%), in concordance with similar TPA typing studies. The majority of the TPA types found were unique per country. New allelic types (7) and profiles (10) were found. The successfully sequenced 23S rRNA loci from 123/137 (90%) samples showed the presence of 79% A2058G and 2% A2059G mutations. CONCLUSIONS: No TEN was found in the samples from different ethnicities residing in Amsterdam, the Netherlands, so no misdiagnoses occurred. Bejel has thus not (yet) spread as a sexually transmitted disease in the Netherlands. The strain diversity found in this study reflects the local male STI clinic population which is a diverse, mixed group.

Testing for changes in biomass dynamics in large-scale forest datasets.

Tropical forest responses to climate and atmospheric change are critical to the future of the global carbon budget. Recent studies have reported increases in estimated above-ground biomass (EAGB) stocks, productivity, and mortality in old-growth tropical forests. These increases could reflect a shift in forest functioning due to global change and/or long-lasting recovery from past disturbance. We introduce a novel approach to disentangle the relative contributions of these mechanisms by decomposing changes in whole-plot biomass fluxes into contributions from changes in the distribution of gap-successional stages and changes in fluxes for a given stage. Using 30 years of forest dynamic data at Barro Colorado Island, Panama, we investigated temporal variation in EAGB fluxes as a function of initial EAGB (EAGBi ) in 10 × 10 m quadrats. Productivity and mortality fluxes both increased strongly with initial quadrat EAGB. The distribution of EAGB (and thus EAGBi ) across quadrats hardly varied over 30 years (and seven censuses). EAGB fluxes as a function of EAGBi varied largely and significantly among census intervals, with notably higher productivity in 1985-1990 associated with recovery from the 1982-1983 El Niño event. Variation in whole-plot fluxes among census intervals was explained overwhelmingly by variation in fluxes as a function of EAGBi , with essentially no contribution from changes in EAGBi distributions. The high observed temporal variation in productivity and mortality suggests that this forest is very sensitive to climate variability. There was no consistent long-term trend in productivity, mortality, or biomass in this forest over 30 years, although the temporal variability in productivity and mortality was so strong that it could well mask a substantial trend. Accurate prediction of future tropical forest carbon budgets will require accounting for disturbance-recovery dynamics and understanding temporal variability in productivity and mortality.

Lightning is a major cause of large tree mortality in a lowland neotropical forest.

The mortality rates of large trees are critical to determining carbon stocks in tropical forests, but the mechanisms of tropical tree mortality remain poorly understood. Lightning strikes thousands of tropical trees every day, but is commonly assumed to be a minor agent of tree mortality in most tropical forests. We use the first systematic quantification of lightning-caused mortality to show that lightning is a major cause of death for the largest trees in an old-growth lowland forest in Panama. A novel lightning strike location system together with field surveys of strike sites revealed that, on average, each strike directly kills 3.5 trees (> 10 cm diameter) and damages 11.4 more. Given lightning frequency data from the Earth Networks Total Lightning Network and historical total tree mortality rates for this site, we conclude that lightning accounts for 40.5% of the mortality of large trees (> 60 cm diameter) in the short term and probably contributes to an additional 9.0% of large tree deaths over the long term. Any changes in cloud-to-ground lightning frequency due to climatic change will alter tree mortality rates; projected 25-50% increases in lightning frequency would increase large tree mortality rates in this forest by 9-18%. The results of this study indicate that lightning plays a critical and previously underestimated role in tropical forest dynamics and carbon cycling.

Role of tree size in moist tropical forest carbon cycling and water deficit responses.

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.

Drought-induced mortality patterns and rapid biomass recovery in a terra firme forest in the Colombian Amazon.

Extreme climatic events affecting the Amazon region are expected to become more frequent under ongoing climate change. In this study, we assessed the responses to the 2010 drought of over 14,000 trees ≥10 cm dbh in a 25 ha lowland forest plot in the Colombian Amazon and how these responses varied among topographically defined habitats, with tree size, and with species wood density. Tree mortality was significantly higher during the 2010-2013 period immediately after the drought than in 2007-2010. The post-drought increase in mortality was stronger for trees located in valleys (+243%) than for those located on slopes (+67%) and ridges (+57%). Tree-based generalized linear mixed models showed a significant negative effect of species wood density on mortality and no effect of tree size. Despite the elevated post-drought mortality, aboveground biomass increased from 2007 to 2013 by 1.62 Mg ha-1  yr-1 (95% CI 0.80-2.43 Mg ha-1  yr-1 ). Biomass change varied among habitats, with no significant increase on the slopes (1.05, 95% CI -0.76 to 2.85 Mg ha-1  yr-1 ), a significant increase in the valleys (1.33, 95% CI 0.37-2.34 Mg ha-1  yr-1 ), and a strong increase on the ridges (2.79, 95% CI 1.20-4.21 Mg ha-1  yr-1 ). These results indicate a high carbon resilience of this forest to the 2010 drought due to habitat-associated and interspecific heterogeneity in responses including directional changes in functional composition driven by enhanced performance of drought-tolerant species that inhabit the drier ridges.

Growth and reproduction respond differently to climate in three Neotropical tree species.

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.

Surviving in a Cosexual World: A Cost-Benefit Analysis of Dioecy in Tropical Trees.

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.

Cascading effects of defaunation on the coexistence of two specialized insect seed predators.

Identification of the mechanisms enabling stable coexistence of species with similar resource requirements is a central challenge in ecology. Such coexistence can be facilitated by species at higher trophic levels through complex multi-trophic interactions, a mechanism that could be compromised by ongoing defaunation. We investigated cascading effects of defaunation on Pachymerus cardo and Speciomerus giganteus, the specialized insect seed predators of the Neotropical palm Attalea butyracea, testing the hypothesis that vertebrate frugivores and granivores facilitate their coexistence. Laboratory experiments showed that the two seed parasitoid species differed strongly in their reproductive ecology. Pachymerus produced many small eggs that it deposited exclusively on the fruit exocarp (exterior). Speciomerus produced few large eggs that it deposited exclusively on the endocarp, which is normally exposed only after a vertebrate handles the fruit. When eggs of the two species were deposited on the same fruit, Pachymerus triumphed only when it had a long head start, and the loser always succumbed to intraguild predation. We collected field data on the fates of 6569 Attalea seeds across sites in central Panama with contrasting degrees of defaunation and wide variation in the abundance of vertebrate frugivores and granivores. Speciomerus dominated where vertebrate communities were intact, whereas Pachymerus dominated in defaunated sites. Variation in the relative abundance of Speciomerus across all 84 sampling sites was strongly positively related to the proportion of seeds attacked by rodents, an indicator of local vertebrate abundance. SYNTHESIS: We show that two species of insect seed predators relying on the same host plant species are niche differentiated in their reproductive strategies such that one species has the advantage when fruits are handled promptly by vertebrates and the other when they are not. Defaunation disrupts this mediating influence of vertebrates and strongly favours one species at the expense of the other, providing a case study of the cascading effects of defaunation and its potential to disrupt coexistence of non-target species, including the hyperdiverse phytophagous insects of tropical forests.

Negative density dependence of seed dispersal and seedling recruitment in a neotropical palm.

Negative density dependence (NDD) of recruitment is pervasive in tropical tree species. We tested the hypotheses that seed dispersal is NDD, due to intraspecific competition for dispersers, and that this contributes to NDD of recruitment. We compared dispersal in the palm Attalea butyracea across a wide range of population density on Barro Colorado Island in Panama and assessed its consequences for seed distributions. We found that frugivore visitation, seed removal and dispersal distance all declined with population density of A. butyracea, demonstrating NDD of seed dispersal due to competition for dispersers. Furthermore, as population density increased, the distances of seeds from the nearest adult decreased, conspecific seed crowding increased and seedling recruitment success decreased, all patterns expected under poorer dispersal. Unexpectedly, however, our analyses showed that NDD of dispersal did not contribute substantially to these changes in the quality of the seed distribution; patterns with population density were dominated by effects due solely to increasing adult and seed density.

Hydrological networks and associated topographic variation as templates for the spatial organization of tropical forest vegetation.

An understanding of the spatial variability in tropical forest structure and biomass, and the mechanisms that underpin this variability, is critical for designing, interpreting, and upscaling field studies for regional carbon inventories. We investigated the spatial structure of tropical forest vegetation and its relationship to the hydrological network and associated topographic structure across spatial scales of 10-1000 m using high-resolution maps of LiDAR-derived mean canopy profile height (MCH) and elevation for 4930 ha of tropical forest in central Panama. MCH was strongly associated with the hydrological network: canopy height was highest in areas of positive convexity (valleys, depressions) close to channels draining 1 ha or more. Average MCH declined strongly with decreasing convexity (transition to ridges, hilltops) and increasing distance from the nearest channel. Spectral analysis, performed with wavelet decomposition, showed that the variance in MCH had fractal similarity at scales of ∼30-600 m, and was strongly associated with variation in elevation, with peak correlations at scales of ∼250 m. Whereas previous studies of topographic correlates of tropical forest structure conducted analyses at just one or a few spatial grains, our study found that correlations were strongly scale-dependent. Multi-scale analyses of correlations of MCH with slope, aspect, curvature, and Laplacian convexity found that MCH was most strongly related to convexity measured at scales of 20-300 m, a topographic variable that is a good proxy for position with respect to the hydrological network. Overall, our results support the idea that, even in these mesic forests, hydrological networks and associated topographical variation serve as templates upon which vegetation is organized over specific ranges of scales. These findings constitute an important step towards a mechanistic understanding of these patterns, and can guide upscaling and downscaling.

Fitting ecological process models to spatial patterns using scalewise variances and moment equations.

Ecological spatial patterns are structured by a multiplicity of processes acting over a wide range of scales. We propose a new method, based on the scalewise variance--that is, the variance as a function of spatial scale, calculated here with wavelet kernel functions--to disentangle the signature of processes that act at different and similar scales on observed spatial patterns. We derive exact and approximate analytical solutions for the expected scalewise variance under different individual-based, spatially explicit models for sessile organisms (e.g., plants), using moment equations. We further determine the probability distribution of independently observed scalewise variances for a given expectation, including complete spatial randomness. Thus, we provide a new analytical test of the null model of spatial randomness to understand at which scales, if any, the variance departs significantly from randomness. We also derive the likelihood function that is needed to estimate parameters of spatial models and their uncertainties from observed patterns. The methods are demonstrated through numerical examples and case studies of four tropical tree species on Barro Colorado Island, Panama. The methods developed here constitute powerful new tools for investigating effects of ecological processes on spatial point patterns and for statistical inference of process models from spatial patterns.

A universal airborne LiDAR approach for tropical forest carbon mapping.

Airborne light detection and ranging (LiDAR) is fast turning the corner from demonstration technology to a key tool for assessing carbon stocks in tropical forests. With its ability to penetrate tropical forest canopies and detect three-dimensional forest structure, LiDAR may prove to be a major component of international strategies to measure and account for carbon emissions from and uptake by tropical forests. To date, however, basic ecological information such as height-diameter allometry and stand-level wood density have not been mechanistically incorporated into methods for mapping forest carbon at regional and global scales. A better incorporation of these structural patterns in forests may reduce the considerable time needed to calibrate airborne data with ground-based forest inventory plots, which presently necessitate exhaustive measurements of tree diameters and heights, as well as tree identifications for wood density estimation. Here, we develop a new approach that can facilitate rapid LiDAR calibration with minimal field data. Throughout four tropical regions (Panama, Peru, Madagascar, and Hawaii), we were able to predict aboveground carbon density estimated in field inventory plots using a single universal LiDAR model (r ( 2 ) = 0.80, RMSE = 27.6 Mg C ha(-1)). This model is comparable in predictive power to locally calibrated models, but relies on limited inputs of basal area and wood density information for a given region, rather than on traditional plot inventories. With this approach, we propose to radically decrease the time required to calibrate airborne LiDAR data and thus increase the output of high-resolution carbon maps, supporting tropical forest conservation and climate mitigation policy.

Linking fruit traits to variation in predispersal vertebrate seed predation, insect seed predation, and pathogen attack.

The importance of vertebrates, invertebrates, and pathogens for plant communities has long been recognized, but their absolute and relative importance in early recruitment of multiple coexisting tropical plant species has not been quantified. Further, little is known about the relationship of fruit traits to seed mortality due to natural enemies in tropical plants. To investigate the influences of vertebrates, invertebrates, and pathogens on reproduction of seven canopy plant species varying in fruit traits, we quantified reductions in fruit development and seed germination due to vertebrates, invertebrates, and fungal pathogens through experimental removal of these enemies using canopy exclosures, insecticide, and fungicide, respectively. We also measured morphological fruit traits hypothesized to mediate interactions of plants with natural enemies of seeds. Vertebrates, invertebrates, and fungi differentially affected predispersal seed mortality depending on the plant species. Fruit morphology explained some variation among species; species with larger fruit and less physical protection surrounding seeds exhibited greater negative effects of fungi on fruit development and germination and experienced reduced seed survival integrated over fruit development and germination in response to vertebrates. Within species, variation in seed size also contributed to variation in natural enemy effects on seed viability. Further, seedling growth was higher for seeds that developed in vertebrate exclosures for Anacardium excelsum and under the fungicide treatment for Castilla elastica, suggesting that predispersal effects of natural enemies may carry through to the seedling stage. This is the first experimental test of the relative effects of vertebrates, invertebrates, and pathogens on seed survival in the canopy. This study motivates further investigation to determine the generality of our results for plant communities. If there is strong variation in natural enemy attack among species related to differences in fruit morphology, then quantification of fruit traits will aid in predicting the outcomes of interactions between plants and their natural enemies. This is particularly important in tropical forests, where high species diversity makes it logistically impossible to study every plant life history stage of every species.