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

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

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

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

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

Humans are increasing nutrient deposition across the globe, and we know little about how these changes influence consumer populations in tropical rainforests. We used a long-term fertilization experiment conducted in a Panamanian forest to explore how nutrient availability and tree traits affect abundance of a higher-level consumer. We added nitrogen, phosphorus and potassium in a factorial design for 18 years. Given that phosphorus often limits ecosystem processes in lowland tropical forests, and added nitrogen reduces insect abundance in our experiment, we first hypothesized that phosphorus addition would increase nest density and nest size of Azteca chartifex ants while nitrogen addition would have the opposite effects. We found 48% lower nest density in the canopy of nitrogen addition plots relative to plots that did not receive nitrogen. Phosphorus addition did not affect nest density or size. These nutrient effects were not diminished by the selectivity of host trees. In general, larger trees held more nests, despite their low frequencies across the forest, while some abundant species (e.g., palms) were rarely used. We further predicted higher nest frequency on trees with extrafloral nectaries, because this ant fuels its large colonies with extrafloral nectar. Despite the non-random distribution of A. chartifex nests, across tree species and nutrient treatments, trees with extrafloral nectaries did not host more nests. Our study suggests that areas of a tropical lowland forest which are not oversaturated with nitrogen, and contain large trees, have higher nest density. This could enable A. chartifex in similar areas to outcompete other ants due to high abundance. [ABSTRACT FROM AUTHOR]

Phenology models developed for temperate and boreal plants predict a single, population‐level first flowering date for each year. These models cannot accommodate species that flower multiple times each year in humid tropical forests nor flowering data with census‐interval rather than daily temporal resolution. Here, we present a new model framework able to predict the timing of multiple annual flowering events from census data. We extend a recent model, which predicted tropical flowering probabilities as discrete events occurring on census dates, by integrating predicted flowering probabilities over all dates between censuses. We evaluate our model against 29 years of daily climate and weekly flowering records for Hybanthus prunifolius (Violaceae) and Handroanthus guayacan (Bignoniaceae) from Barro Colorado Island, Panama. Previous experiments demonstrate that both species flower shortly after a heavy, dry‐season rain interrupts an extended dry period. Our model captures this sequence of rainfall events. Best‐fit model parameters are consistent with previous experimental results. This match suggests the new model framework will provide novel insights for other humid tropical forest species. We present a new model framework to evaluate hypotheses concerning environmental cues for phenology using observational records. We validate the approach using 29 years of daily climate records and weekly flowering observations for two tropical tree species whose flowering cues are known from prior experiments. The new model framework successfully identified the experimentally determined flowering cues promising new insights for the vast majority of species whose phenology has not been manipulated experimentally. [ABSTRACT FROM AUTHOR]

Numerous Neotropical rainforest species are distributed in both Amazonia and Central America, reflecting a rich history of biotic interchange between regions. However, some plant lineages are endemic to one region, due in part to the dispersal barrier posed by the Northern Andean Cordilleras and adjacent savannas. To investigate the role of biogeographic filtering across the northern Andes in regional community assembly, we examined environmental tolerances, functional traits, and biogeographic distributions of >1000 woody plant species (trees, shrubs, lianas) locally co‐occurring in forest plots in lowland Panama (542 species) and Amazonian Ecuador (667 species). High regional abundance was strongly predictive of the probability of being geographically widespread (i.e. present on both sides of the Andes). However, we also found that species with broad environmental tolerances (those that are able to inhabit high elevations and areas of low mean annual precipitation) were more likely to have a cross‐Andean distribution even after accounting for regional abundance, suggesting that biogeographic filtering for these traits has mediated cross‐Andean dispersal. Regional abundance and environmental tolerances were additionally associated with a suite of life‐history traits related to high dispersal–colonization ability, but most traits reflecting dispersal–colonization ability were not predictive of biogeographic distribution. Our results highlight how the process of biogeographic filtering, based primarily on environmental tolerances, has mediated regional‐scale floristic assembly of Neotropical rainforests. The impacts of this process, which we term filter‐dispersal assembly, are likely to be especially important to forests in Central America, where biotic interchange with Amazonia has heavily influenced regional community composition. [ABSTRACT FROM AUTHOR]

With rising atmospheric CO2 concentrations, strategies for greater carbon sequestration and long‐term storage are urgently needed. Current reforestation schemes typically use fast‐growing tree species with low wood density, limiting the amount of carbon stored. In most tropical forests, there are native species with relatively fast growth rates, high wood density and low mortality rates that also achieve large adult size—'exceptional species'.We present a simulated assessment of the carbon sequestration potential using these exceptional species in assisted secondary succession in the Neotropics.In our example, the selected exceptional species could add up to 16.2 Mg C/ha of sequestered carbon in secondary wet forests at maturity.Synthesis and applications. By increasing the representation of exceptional species in reforestation efforts, we could potentially boost long‐term carbon storage in reforestation projects focussed on carbon sequestration. However, our approach is not a substitute for protecting existing old primary forests and reforested lands plus reducing deforestation and emissions. [ABSTRACT FROM AUTHOR]

Abstract: We present a meta‐analysis of plant responses to fertilization experiments conducted in lowland, species‐rich, tropical forests. We also update a key result and present the first species‐level analyses of tree growth rates for a 15‐yr factorial nitrogen (N), phosphorus (P), and potassium (K) experiment conducted in central Panama. The update concerns community‐level tree growth rates, which responded significantly to the addition of N and K together after 10 yr of fertilization but not after 15 yr. Our experimental soils are infertile for the region, and species whose regional distributions are strongly associated with low soil P availability dominate the local tree flora. Under these circumstances, we expect muted responses to fertilization, and we predicted species associated with low‐P soils would respond most slowly. The data did not support this prediction, species‐level tree growth responses to P addition were unrelated to species‐level soil P associations. The meta‐analysis demonstrated that nutrient limitation is widespread in lowland tropical forests and evaluated two directional hypotheses concerning plant responses to N addition and to P addition. The meta‐analysis supported the hypothesis that tree (or biomass) growth rate responses to fertilization are weaker in old growth forests and stronger in secondary forests, where rapid biomass accumulation provides a nutrient sink. The meta‐analysis found no support for the long‐standing hypothesis that plant responses are stronger for P addition and weaker for N addition. We do not advocate discarding the latter hypothesis. There are only 14 fertilization experiments from lowland, species‐rich, tropical forests, 13 of the 14 experiments added nutrients for five or fewer years, and responses vary widely among experiments. Potential fertilization responses should be muted when the species present are well adapted to nutrient‐poor soils, as is the case in our experiment, and when pest pressure increases with fertilization, as it does in our experiment. The statistical power and especially the duration of fertilization experiments conducted in old growth, tropical forests might be insufficient to detect the slow, modest growth responses that are to be expected. [ABSTRACT FROM AUTHOR]

Producing and retaining leaves underlie the performance and survivorship of seedlings in deeply shaded tropical forests. These habitats are characterized by conditions ideal for foliar bacteria, which can be potent plant pathogens. Leaf production, retention and susceptibility to enemies may ultimately depend upon interactions among soil nutrients and foliar microbes, yet this has never been tested., We experimentally evaluated the degree that foliar bacteria and soil resource supply mediate leaf dynamics for five common tree species (five different families) in a Panamanian forest. We reduced foliar bacteria with antibiotics for 29 months and measured leaf production, retention and damage for seedlings nested within a replicated 15-yr factorial nutrient enrichment experiment (nitrogen, N; phosphorus, P; potassium, K)., Our results demonstrate that when we applied antibiotics, soil nutrients - particularly N - always regulated seedling leaf production (and to a lesser extent herbivore damage) for all five tree species. In addition, it was common for two macronutrients together to negate or completely reverse the impact of applying either one alone., Our findings of frequent plant-microbe-nutrient interactions are novel and suggest that these interactions may reinforce plant species-environment associations, thereby creating a fairly cryptic and fine-scale dimension of niche differentiation for coexisting tree species. [ABSTRACT FROM AUTHOR]

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

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. [ABSTRACT FROM AUTHOR]

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

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

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-m2 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. [ABSTRACT FROM AUTHOR]

Seed size and toughness affect seed predators, and size-dependent investment in mechanical defence could affect relationships between seed size and predation. We tested how seed toughness and mechanical defence traits (tissue density and protective tissue content) are related to seed size among tropical forest species. Absolute toughness increased with seed size. However, smaller seeds had higher specific toughness both within and among species, with the smallest seeds requiring over 2000 times more energy per gram to break than the largest seeds. Investment in mechanical defence traits varied widely but independently of the toughness-mass allometry. Instead, a physical scaling relationship confers a toughness advantage on small seeds independent of selection on defence traits and without a direct cost. This scaling relationship may contribute to seed size diversity by decreasing fitness differences among large and small seeds. Allometric scaling of toughness reconciles predictions and conflicting empirical relationships between seed size and predation. [ABSTRACT FROM AUTHOR]

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

The importance of lianas through time and their effect on tree reproduction are evaluated for the first time in a Southeast Asian Dipterocarp forest. We quantified flower and seed production by lianas and trees for 13 years, assessed liana loads in the crowns of all trees larger than 30 cm in diameter at breast height (1.3 m) in 2002 and 2014, and assessed levels of reproduction for the same trees during a strong general flowering event in 2014 for the 50-ha forest dynamics plot at the Pasoh Forest Reserve, Malaysia. General flowering refers to synchronous reproduction by hundreds of plant species at irregular, multiyear intervals and only occurs in Southeast Asian Dipterocarp forests. Overall, lianas were present in 50% of tree crowns and comprised 31% of flower production and 46% of seed production. Lianas reduced growth, survival, and reproduction by their host trees. Lianas were less frequent in canopyemergent trees, Dipterocarps comprised a disproportionately large proportion of canopy emergents, and, as a consequence, lianas were less frequent in Dipterocarps than in trees from other plant families. Lianas infested the crowns of significantly fewer trees in 2014 (47.9%) than in 2002 (52.3%); however, the decrease was restricted to trees with the lightest liana loads and sample sizes and statistical power were enormous. Lianas comprised a stable proportion of flower production and a highly variable proportion of seed production from 2002 through 2013. We conclude lianas have a huge impact on trees in this forest and were a stable component of the forest between 2002 and 2014. The emergent habit and associated ability to avoid lianas might contribute to the success of the Dipterocarpaceae. [ABSTRACT FROM AUTHOR]

Questions remain as to which soil nutrients limit primary production in tropical forests. Phosphorus (P) has long been considered the primary limiting element in lowland forests, but recent evidence demonstrates substantial heterogeneity in response to nutrient addition, highlighting a need to understand and diagnose nutrient limitation across diverse forests. Fine-root characteristics including their abundance, functional traits, and mycorrhizal symbionts can be highly responsive to changes in soil nutrients and may help to diagnose nutrient limitation. Here, we document the response of fine roots to long-term nitrogen (N), P, and potassium (K) fertilization in a lowland forest in Panama. Because this experiment has demonstrated that N and K together limit tree growth and P limits fine litter production, we hypothesized that fine roots would also respond to nutrient addition. Specifically we hypothesized that N, P, and K addition would reduce the biomass, diameter, tissue density, and mycorrhizal colonization of fine roots, and increase nutrient concentration in root tissue. Most morphological root traits responded to the single addition of K and the paired addition of N and P, with the greatest response to all three nutrients combined. The addition of N, P, and K together reduced fine-root biomass, length, and tissue density, and increased specific root length, whereas root diameter remained unchanged. Nitrogen addition did not alter root N concentration, but P and K addition increased root P and K concentration, respectively. Mycorrhizal colonization of fine roots declined with N, increased with P, and was unresponsive to K addition. Although plant species composition remains unchanged after 14 years of fertilization, fine-root characteristics responded to N, P, and K addition, providing some of the strongest stand-level responses in this experiment. Multiple soil nutrients regulate fine-root abundance, morphological and chemical traits, and their association with mycorrhizal fungi in a species-rich lowland tropical forest. [ABSTRACT FROM AUTHOR]

Leaf defences vary widely among tree species, affecting rates of herbivory, survival and reproduction., Two contrasting hypotheses account for variation in leaf defences among species. The first predicts that a slow life history, which is characteristic of larger seeded species adapted to resource-limited environments, is associated with well-defended leaves. The second, apparency theory, predicts that elevated leaf defences are necessitated for species that are more detectable to herbivores., Here we use comparative methods and a global data set to test (i) the relationship between seed size and leaf defences and (ii) the relationship between clumping (spatial apparency) and leaf defences., We found that seed size was positively related to leaf fracture toughness, but not phenolics or tannin concentration and that spatial aggregation was unrelated to leaf defences., Synthesis. Our results suggest that larger seed size and increased leaf toughness are correlated as part of a trait syndrome associated with a slow, resource-limited life history, not clumped dispersion and increased spatial apparency. [ABSTRACT FROM AUTHOR]

Defaunation alters trophic interactions between plants and vertebrates, which may disrupt trophic cascades, thereby favoring a subset of plant species and reducing diversity. If particular functional traits characterize the favored plant species, then defaunation may alter community-wide patterns of functional trait composition. Changes in plant functional traits occurring with defaunation may help identify the species interactions affected by defaunation and the potential for other cascading effects of defaunation. We tested the hypotheses that defaunation would (1) disrupt seed dispersal, thereby favoring species whose dispersal agents are not affected (e.g., small birds, bats, and abiotic agents), (2) reduce seed predation, thereby favoring larger-seeded species, and (3) reduce herbivory, thereby favoring species with lower leaf mass per area (LMA), leaf toughness, and wood density. We examined how these six traits responded to vertebrate defaunation caused by hunters or by experimental exclosures among more than 30 000 woody seedlings in a lowland tropical moist forest. Exclosures reduced terrestrial frugivores, granivores, and herbivores, while hunters also reduced volant and arboreal frugivores and granivores. The comparison of exclosures and hunting allowed us to parse the impacts of arboreal and volant species (reduced by hunters only) and terrestrial species (reduced by both hunters and exclosures). The loss of terrestrial vertebrates alone had limited effects on plant trait composition. The additional loss of volant and arboreal vertebrates caused significant shifts in plant species composition towards communities with more species dispersed abiotically, including lianas and low wood-density tree species, and fewer species dispersed by large vertebrates. In contrast to previous studies, community seed mass did not decline significantly in hunted sites. Our exclosure results suggest this is because reducing seed predators disproportionately benefits large-seeded species, partially compensating for the reduction of seed dispersers at hunted sites. Our results demonstrate that decreased seed dispersal and seed predation are important determinants of seedling community compositional change as a consequence of defaunation. Defaunation may also negatively impact carbon storage in tropical forests, by favoring lianas and low wood density tree species. [ABSTRACT FROM AUTHOR]

Lianas are a prominent growth form in tropical forests, and there is compelling evidence that they are increasing in abundance throughout the Neotropics. While recent evidence shows that soil resources limit tree growth even in deep shade, the degree to which soil resources limit lianas in forest understories, where they coexist with trees for decades, remains unknown. Regardless, the physiological underpinnings of soil resource limitation in deeply shaded tropical habitats remain largely unexplored for either trees or lianas. Theory predicts that lianas should be more limited by soil resources than trees because they occupy the quick-return end of the ''leaf economic spectrum,'' characterized by high rates of photosynthesis, high specific leaf area, short leaf life span, affinity to high-nutrient sites, and greater foliar nutrient concentrations. To address these issues, we asked whether soil resources (nitrogen, phosphorus, and potassium), alone or in combination, applied experimentally for more than a decade would cause significant changes in the morphology or physiology of tree and liana seedlings in a lowland tropical forest. We found evidence for the first time that phosphorus limits the photosynthetic performance of both trees and lianas in deeply shaded understory habitats. More importantly, lianas always showed significantly greater photosynthetic capacity, quenching, and saturating light levels compared to trees across all treatments. We found little evidence for nutrient 3 growth form interactions, indicating that lianas were not disproportionately favored in nutrient-rich habitats. Tree and liana seedlings differed markedly for six key morphological traits, demonstrating that architectural differences occurred very early in ontogeny prior to lianas finding a trellis (all seedlings were self-supporting). Overall, our results do not support nutrient loading as a mechanism of increasing liana abundance in the Neotropics. Rather, our finding that lianas always outperform trees, in terms of photosynthetic processes and under contrasting rates of resource supply of macronutrients, will allow lianas to increase in abundance if disturbance and tree turnover rates are increasing in Neotropical forests as has been suggested. [ABSTRACT FROM AUTHOR]

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

A complex set of interactions among neighbors influences plant performance and community structure. Understanding their joint operation requires extensive information on species characteristics and individual performance. We evaluated first-year survival of 35 719 tropical forest seedlings of 222 species and 15 annual cohorts relative to the density of conspecific and heterospecific neighbors and the phylogenetic similarity of heterospecific neighbors. Neighbors were from two size classes, and size asymmetric interactions provided insight into likely mechanisms. Large heterospecific and conspecific neighbors reduced seedling survival equally, suggesting resource competition rather than host-specific enemies as a mechanism. In contrast, much stronger negative conspecific effects were associated with seedling neighbors capable of limited resource uptake, suggesting shared pests rather than competition as the mechanism. Survival improved, however, near phylogenetically similar heterospecific neighbors, suggesting habitat associations shared among closely related species affect spatial patterns of performance. Improved performance near phylogenetically similar neighbors is an emerging pattern in the handful of similar studies. [ABSTRACT FROM AUTHOR]

The concentration, stoichiometry and resorption of nitrogen ( N) and phosphorus ( P) in plant leaves are often used as proxies of the availability of these growth-limiting nutrients, but the responses of these metrics to changes in nutrient availability remain largely untested for tropical forest trees., We evaluated changes in N and P concentrations, N/ P ratios and resorption for four common tree species after 13 years of factorial N and P additions in a lowland tropical forest in Panama., Chronic P addition increased foliar P concentrations, decreased P resorption proficiency and decreased N/ P ratios in three locally common eudicot tree species ( Alseis blackiana, Heisteria concinna, Tetragastris panamensis). The increase in foliar P involved similar proportional increases in organic and inorganic P in two species and a disproportionately large increase in inorganic P in A. blackiana., Nitrogen addition did not alter foliar N concentrations in any species, but did decrease N resorption proficiency in H. concinna., A fourth species, the palm Oenocarpus mapora, demonstrated remarkably static foliar nutrient concentrations, responding only with a marginal decrease in P resorption proficiency under N plus P co-addition., Synthesis. Collectively, these results suggest that adjustment of N/ P ratios can be expected in eudicots exposed to elevated P, but foliar N appears to already be at optimal levels in these lowland rain forest tree species. The complexity of species-specific responses to altered nutrient availability highlights the difficulty in predicting future responses of tropical forest trees to a changing world. [ABSTRACT FROM AUTHOR]

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

1. A small number of species-rich plant genera make a substantial contribution to the α diversity of tropical forests. These 'species swarms' (Gentry 1982) challenge a view of community assembly that maintains that niche overlap, and hence, the likelihood of competitive exclusion should increase with phylogenetic affinity. Related species may, however, occupy different microhabitats within a forest stand or, alternatively, differ ecologically in ways that allow them to coexist syntopically. 2. To explore the relationship between phylogenetic history and niche differentiation among sympatric, congeneric species, we performed phylogenetic analyses of microhabitat preferences, photosynthetic and hydraulic traits, and experimental responses to light and water availability for 20 species of Psychotria (Rubiaceae) from Barro Colorado Island (BCI), Panama. 3. The Psychotria present in small (3-m radius), circular plots were more closely related than expected by chance. Photosynthetic and hydraulic traits were both associated with species habitat distributions, but only hydraulic traits were conserved phylogenetically. Experimental responses to water availability were also conserved phylogenetically. Functional trait analyses revealed little or no evidence of niche partitioning within sites. 4. We conclude that Psychotria species' responses to both light and moisture availability determine their microhabitat distributions on BCI and that evolutionarily conserved hydraulic traits lead to phylogenetic clustering of co-occurring species. 5. Synthesis. The evolutionary conservation of hydraulic traits related to soil moisture tolerance largely explains phylogenetic clustering in the local assembly of the hyperdiverse genus Psychotria. We suggest that close relatives are unlikely to exclude one another from shared habitats because resource availability is determined largely by asymmetric competition with the overstorey, rather than by competition with neighbouring understorey plants. In light of the recent biogeographic admixture in central Panama, the phylogenetic niche conservatism exhibited by Psychotria on BCI raises the possibility of an association between local microhabitats and the ancestral climatic regimes under which major Psychotria lineages evolved before arriving in sympatry. [ABSTRACT FROM AUTHOR]

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

Summary 1. Nutrients are a critical resource for plant growth, but the elements limiting growth in tropical forests have rarely been determined. 2. We investigated the influence of nitrogen (N), phosphorus (P), potassium (K) and micronutrients on seedling biomass and nutrient allocation in a factorial nutrient fertilization experiment in lowland tropical forest at the Barro Colorado Nature Monument, Panama. We also measured 8 years of herbivory and growth for 1800 seedlings. We sought to determine the identity of limiting elements and possible nutrient interactions. 3. The five study species were Alseis blackiana, Desmopsis panamensis, Heisteria concinna, Sorocea affinis and Tetragastris panamensis. Plants grew in deeply shaded understorey with a mean canopy openness of 4.9% (±0.7%; 1 SE). 4. Tissue N concentration increased by 11% with N addition. Tissue P concentration increased by 16% with P addition. Tissue K increased by 4% with K addition. K addition reduced root-to-shoot biomass ratio. There was no significant effect of fertilization on specific leaf area or leaf area ratio. 5. The proportion of leaves damaged and the mean level of damage by herbivory increased with P and K addition and showed a significant P × K interaction. 6.7ensp;Across all species and years, relative growth rate of height increased with K addition and with N and P in combination. Relative growth rate of leaf count trended 8.5% higher with K addition ( P = 0.076). 7. We also added micronutrients in a parallel experiment. There was no effect of micronutrient addition on any seedling parameter. 8. Synthesis. K addition affected seedlings by enhancing tissue nutrient concentration, increasing herbivory, reducing root-to-shoot biomass ratio and increasing height growth. Additional effects of N or P on tissue chemistry, herbivory and growth offer support for the multiple limiting resources hypothesis. Our results suggest that seedling growth is limited by nutrients, especially K, even under highly shaded conditions in this lowland tropical forest. [ABSTRACT FROM AUTHOR]

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

1. Lianas compete intensely with trees, but few studies have examined long-term effects of liana infestation on tree growth and mortality. We quantified the effects of lianas in tree crowns ( n = 2907) and rooted within 2 m of trees ( n = 1086) on growth and mortality of 30 tree species from 1995 to 2005 on Barro Colorado Island (BCI), Panama, documented liana infestation in tree crowns in 1996 and 2007 to determine the dynamics of liana infestation, and quantified liana infestation in the crowns of 3231 additional canopy trees (d.b.h. ≥20 cm) in 2007 to compare with the same metric determined by previous studies in 1967 and 1980. 2. Severe liana infestation increased tree mortality: 21% of liana-free trees in 1996 had died by 2007, whereas 42% of trees with more than 75% of the crown infested by lianas in 1996 had died by 2007. 3. Liana infestation of tree crowns significantly reduced tree growth, particularly on sun-exposed trees. The proximity of rooted lianas significantly reduced the growth of shaded trees. 4. Liana infestation was dynamic: 10.9% of trees with severe liana infestation in their crowns in 1996 had shed all of their lianas by 2007 and 5.3% of trees with no lianas in their crown in 1996 had severe liana infestation in 2007. 5. Liana infestation was common: lianas were present in 53% of trees of the 30 focal species. Including lianas rooted within 2 m of the tree increased this percentage to 78%. Using both above- and below-ground measures may provide a better estimate of liana competition than either measure alone. 6. Liana infestation is increasing on BCI. Lianas were present in the crowns of 73.6% of canopy trees (d.b.h. ≥20 cm). Liana canopy infestation was 57% higher than in 1980 and 65% higher than in 1967, which is consistent with reported increases in liana abundance, biomass, and leaf and flower production. 7. Synthesis. We used one of the largest studies ever conducted on lianas to confirm the negative effects of lianas on tree growth and survival over 10 years. Liana infestation of trees was widespread, dynamic and increasing on BCI. [ABSTRACT FROM AUTHOR]

Five anthropogenic drivers–land use change, wood extraction, hunting, atmospheric change, climate change–will largely determine the future of tropical forests. The geographic scope and intensity of these five drivers are in flux. Contemporary land use change includes deforestation (∼64,000 km2 yr−1 for the entire tropical forest biome) and natural forests regenerating on abandoned land (∼21,500 km2 yr−1 with just 29% of the biome evaluated). Commercial logging is shifting rapidly from Southeast Asia to Africa and South America, but local fuelwood consumption continues to constitute 71% of all wood production. Pantropical rates of net deforestation are declining even as secondary and logged forests increasingly replace old-growth forests. Hunters reduce frugivore, granivore and browser abundances in most forests. This alters seed dispersal, seed and seedling survival, and hence the species composition and spatial template of plant regeneration. Tropical governments have responded to these local threats by protecting 7% of all land for the strict conservation of nature—a commitment that is only matched poleward of 40°S and 70°N. Protected status often fails to stop hunters and is impotent against atmospheric and climate change. There are increasing reports of stark changes in the structure and dynamics of protected tropical forests. Four broad classes of mechanisms might contribute to these changes. Predictions are developed to distinguish among these mechanisms. [ABSTRACT FROM AUTHOR]

We reviewed the evidence on the extent and efficacy of conservation of tropical forest biodiversity for each of the classes of conservation action defined by the new International Union for Conservation of Nature (IUCN) classification. Protected areas are the most tested conservation approach, and a number of studies show they are generally effective in slowing deforestation. There is some documentation of the extent of sustainable timber management in tropical forest, but little information on other landscape-conservation tactics. The extent and effectiveness of ex situ species conservation is quite well known. Forty-one tropical-forest species now survive only in captivity. Other single-species conservation actions are not as well documented. The potential of policy mechanisms, such as international conventions and provision of funds, to slow extinctions in tropical forests is considerable, but the effects of policy are difficult to measure. Finally, interventions to promote tropical conservation by supporting education and livelihoods, providing incentives, and furthering capacity building are all thought to be important, but their extent and effectiveness remain poorly known. For birds, the best studied taxon, the sum of such conservation actions has averted one-fifth of the extinctions that would otherwise have occurred over the last century. Clearly, tropical forest conservation works, but more is needed, as is critical assessment of what works in what circumstances, if mass extinction is to be averted. [ABSTRACT FROM AUTHOR]

Modern global temperature and land cover and projected future temperatures suggest that tropical forest species will be particularly sensitive to global warming. Given a moderate greenhouse gas emissions scenario, fully 75% of the tropical forests present in 2000 will experience mean annual temperatures in 2100 that are greater than the highest mean annual temperature that supports closed-canopy forest today. Temperature-sensitive species might extend their ranges to cool refuges, defined here as areas where temperatures projected for 2100 match 1960s temperatures in the modern range. Distances to such cool refuges are greatest for equatorial species and are particularly large for key tropical forest areas including the Amazon and Congo River Basins, West Africa, and the upper elevations of many tropical mountains. In sum, tropical species are likely to be particularly sensitive to global warming because they are adapted to limited geographic and seasonal variation in temperature, already lived at or near the highest temperatures on Earth before global warming began, and are often isolated from cool refuges. To illustrate these three points, we examined the distributions and habitat associations of all extant mammal species. The distance to the nearest cool refuge exceeded 1000 km for more than 20% of the tropical and less than 4% of the extratropical species with small ranges. The biological impact of global warming is likely to be as severe in the tropics as at temperate and boreal latitudes. [ABSTRACT FROM AUTHOR]

The article discusses various reports published within the issue, including one by Joseph Wright and colleagues on the fate of tropical biodiversity, one by Greg Asner and colleagues on the changes in primary, secondary, and selectively logged forests across the tropical regions in the world, and one by Robin Chazdon and colleagues on the persistence degree of tropical forest species in regeneration forests.

Traits associated with seed dispersal vary tremendously among sympatric wind-dispersed plants. We used two contrasting tropical tree species, seed traps, micrometeorology, and a mechanistic model to evaluate how variation in four key traits affects seed dispersal by wind. The conceptual framework of movement ecology, wherein external factors (wind) interact with internal factors (plant traits) that enable movement and determine when and where movement occurs, fully captures the variable inputs and outputs of wind dispersal models and informs their interpretation. We used model calculations to evaluate the spatial pattern of dispersed seeds for the 16 factorial combinations of four traits. The study species differed dramatically in traits related to the timing of seed release, and a strong species by season interaction affected most aspects of the spatial pattern of dispersed seeds. A rich interplay among plant traits and seasonal differences in atmospheric conditions caused this interaction. Several of the same plant traits are crucial for both seed dispersal and other aspects of life history variation. Observed traits that limit dispersal are likely to be constrained by their life history consequences. [ABSTRACT FROM AUTHOR]

1. We investigated the relationships of seed size, dispersal mode and other species characteristics to interspecific variation in mean primary seed dispersal distances, mean annual seed production per unit basal area, and clumping of seed deposition among 41 tropical tree species on Barro Colorado Island, Panama. 2. A hierarchical Bayesian model incorporating interannual variation in seed production was used to estimate seed dispersal, seed production, and clumping of seed rain for each species from 19 years of data for 188 seed traps on a 50-ha plot in which all adult trees were censused every 5 years. 3. Seed dispersal was modelled as a two-dimensional Student's T distribution with the degrees of freedom parameter fixed at 3, interannual variation in seed production per basal area was modelled as a lognormal, and the clumping of seed rain around its expected value was modelled as a negative binomial distribution. 4. There was wide variation in seed dispersal distances among species sharing the same mode of seed dispersal. Seed dispersal mode did not explain significant variation in seed dispersal distances, but did explain significant variation in clumping: animal-dispersed species showed higher clumping of seed deposition. 5. Among nine wind-dispersed species, the combination of diaspore terminal velocity, tree height and wind speed in the season of peak dispersal explained 40% of variation in dispersal distances. Among 31 animal-dispersed species, 20% of interspecific variation in dispersal distances was explained by seed mass (a negative effect) and tree height (a positive effect). 6. Among all species, seed mass, tree height and dispersal syndrome explained 28% of the variation in mean dispersal distance and seed mass alone explained 45% of the variation in estimated seed production per basal area. 7. Synthesis. There is wide variation in patterns of primary seed rain among tropical tree species. Substantial proportions of interspecific variation in seed production, seed dispersal distances, and clumping of seed deposition are explained by relatively easily measured plant traits, especially dispersal mode, seed mass, and tree height. This provides hope for trait-based generalization and modelling of seed dispersal in tropical forests. [ABSTRACT FROM AUTHOR]

Heterostyly is a genetic polymorphism in which plant populations are composed of two or more morphs that differ in stigma and anther heights. The polymorphism promotes intermorph pollen transfer, thus outcrossing. Heterostyly has been reported in 28 angiosperm families and is frequently lost in heterostylous lineages. To assess ecological factors related to shifts from heterostyly to monomorphy, we examined the reproductive ecology of heterostylous tropical shrubs of Psychotria. Among 21 species at Barro Colorado Island and the nearby Parque Nacional Soberania, Panama, 14 species were heterostylous while seven were monomorphic. A molecular phylogeny and the existence elsewhere of heterostylous populations indicated that the breakdown of heterostyly had occurred independently. Heterostylous and monomorphic species were visited by the same bee species, although visit frequencies were lower in monomorphic species. Monomorphic species had significantly lower population density and greater fruit set than did heterostylous species. Autonomous autogamy made a large contribution to fruit set in monomorphic species and was only rarely observed in heterostylous species. The results indicate monomorphic and heterostylous species produce more seeds through selfing and outcrossing, respectively. The limitation of outcrossing as a result of low population density may be related to the breakdown of heterostyly and the evolution of selfing. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 93, 125–134. [ABSTRACT FROM AUTHOR]

It has been hypothesized that tropical trees partition forest light environments through a life history trade-off between juvenile growth and survival; however, the generality of this trade-off across life stages and functional groups has been questioned. We quantified trade-offs between growth and survival for trees and lianas on Barro Colorado Island (BCI), Panama using first-year seedlings of 22 liana and 31 tree species and saplings (10mm < dbh < 39 mm) of 30 tree species. Lianas showed trade-offs similar to those of trees, with both groups exhibiting broadly overlapping ranges in survival and relative growth rates as seedlings. Life history strategies at the seedling stage were highly correlated with those at the sapling stage among tree species, with all species showing an increase in survival with size. Only one of 30 tree species demonstrated a statistically significant ontogenetic shift, having a relatively lower survival rate at the sapling stage than expected. Our results indicate that similar life history trade-offs apply across two functional groups (lianas and trees), and that life history strategies are largely conserved across seedling and sapling life-stages for most tropical tree species. [ABSTRACT FROM AUTHOR]

Understanding and mitigating the impact of an ever-increasing population and global economic activity on tropical forests is one of the great challenges currently facing biologists, conservationists and policy makers. Tropical forests currently face obvious regional changes, both negative and positive, and uncertain global changes. Although deforestation rates have increased to unprecedented levels, natural secondary succession has reclaimed approximately 15% of the area deforested during the 1990s. Governments have also protected 18% of the remaining tropical moist forest; however, unsustainable hunting continues to threaten many keystone mammal and bird species. The structure and dynamics of old-growth forests appear to be rapidly changing, suggesting that there is a pantropical response to global anthropogenic forcing, although the evidence comes almost exclusively from censuses of tree plots and is controversial. Here, I address ongoing anthropogenic change in tropical forests and suggest how these forests might respond to increasing anthropogenic pressure. [Copyright &y& Elsevier]

1 The role of seed limitation in tropical forests remains uncertain owing to the scarcity of experimental evidence. We performed seed addition experiments to assess seed limitation for 32 shade-tolerant tropical forest species and monitored the natural seed rain of 25 of these species for 17 years. 2 One, two or five seeds were sown into 0.0079-m2 plots for large- ( n = 5 species), medium- ( n = 5) and small-seeded species ( n = 22), respectively. The experiment was replicated at 69 sites, placed in groups of three at 23 locations. Seedling establishment was evaluated after 1 and 2 years in paired seed addition and control plots. Natural seedling emergence and understorey plant density were also measured. 3 Median natural seed rain was 0.31 seeds m−2 year−1 per focal species. 4 Seed addition enhanced seedling establishment in 31 and 26 of the 32 species after 1 and 2 years, respectively. Mean number of focal species’ seedlings after 2 years was 0.002 seedlings in control plots and 0.12, 0.37 and 0.60 seedlings in seed addition plots for large-, medium- and small-seeded species, respectively. 5 A 25 seeds added treatment increased seedling establishment by ≥ 2.0-fold over the five seeds added treatment after 2 years. 6 Community-wide recruitment and understorey plant density were strongly seed-limited. The natural density of understorey plants averaged 12 plants m−2 and was significantly less than for seedlings of the single focal species in plots with ≥ 2 seeds added 2 years earlier. 7 The number of established seedlings per seed added was independent of seed size. 8 Treatment (adding zero or five seeds), species identity and location all affected seedling establishment for the 11 small-seeded species represented at all sites, with treatment and its interactions accounting for 86% of the explained variation. 9 Our results suggest that seed limitation plays a dominant role in seedling recruitment and understorey plant community assembly in tropical forests. Although strong seed limitation may set the stage for species-neutral community assembly, the species differences in seedling establishment rate and its spatial variation demonstrate an important role for species-specific processes. [ABSTRACT FROM AUTHOR]

An economy of scale may lead to selection to increase interannual variation in seed production when the per seed probability of seedling establishment increases with seed production. Variable annual seedfall will, however, reduce this probability when post- dispersal seed fate is negatively density dependent on the local density of seeds, and seed dispersal and density dependence act identically across years. Intuitively, more variable annual seedfall causes the representative seed to experience a greater density of conspecific seeds and suffer greater density-dependent effects. This handicap must be overcome for the per seed probability of recruitment to be greater in years with greater seed production. We quantified spatial and annual variation in seedfall and seedling recruitment, evaluated density dependence and economies of scale during the seed-to-seedling transition, and investigated the synergistic consequences of density dependence and variable annual seed- fall for seedling recruitment on Barro Colorado Island (BCI), Panama. Weekly censuses of 200 0.5-m² seed traps documented seedfall for 15 years and 108 plant species. Annual censuses of 600 1-m² seedling plots documented recruitment for nine years and 32 species. The density of seedling recruits tended to increase with the density of seeds; however, the per seed probability of recruitment invariably decreased with seedfall density. Negative density dependence characterized the seed-to-seedling transition. Observed levels of spatial and interannual variation in seedfall density would reduce long-term recruitment by up to 28% if negatively density-dependent survival acted identically across years; however, the strength of negative density dependence varied significantly among years for 12 of 32 species. Negative density dependence occurred in all years for these species but was significantly weaker during the one or two years of greatest seedfall than during the remaining years of lower seedfall. The per seed probability of recruitment increased significantly with annual seedfall for eight of these species. These eight species realized postdispersal economies of scale despite the reduction in long-term recruitment expected from the synergism between variable annual seed production and negatively density-dependent seed fate. [ABSTRACT FROM AUTHOR]

Climatic conditions should not hinder nutrient release from decomposing leaf-litter (mineralization) in the humid tropics, even though many tropical forests experience drought lasting from several weeks to months. We used a dry-season irrigation experiment to examine the effect of seasonal drought on nutrient concentrations in leaf-fall and in decomposing leaf-litter. In the experiment, soil in two 2.25-ha plots of old-growth lowland moist forest on Barro Colorado Island, Republic of Panama, was watered to maintain soil water potential at or above field capacity throughout the 4-month dry season. Wet-season leaf-fall had greater concentrations of nitrogen (N, 13.5 mg g−1) and calcium (Ca, 15.6 mg g−1) and lower concentrations of sulfur (S, 2.51 mg g−1) and potassium (K, 3.03 mg g−1) than dry-season leaf-fall (N = 11.6 mg g−1, Ca = 13.6 mg g−1, S = 2.98 mg g−1, K = 5.70 mg g−1). Irrigation did not affect nutrient concentrations or nutrient return from forest trees to the forest floor annually (N = 18 g m−2, phosphorus (P) = 1.06 g m−2, S = 3.5 g m−2, Ca = 18.9 g m−2, magnesium = 6.5 g m−2, K = 5.7 g m−2). Nutrient mineralization rates were much greater during the wet season than the dry season, except for K, which did not vary seasonally. Nutrient residence times in forest-floor material were longer in control plots than in irrigated plots, with values approximately equal to that for organic matter (210 in control plots vs 160 in irrigated plots). Calcium had the longest residence time. Forest-floor material collected at the transition between seasons and incubated with or without leaching in the laboratory did not display large pulses in nutrient availability. Rather, microorganisms immobilized nutrients primarily during the wet season, unlike observations in tropical forests with longer dry seasons. Large amounts of P moved among different pools in forest-floor material, apparently mediated by microorganisms. Arylsulfatase and phosphatase enzymes, which mineralize organically bound nutrients, had high activity throughout the dry season. Low soil moisture levels do not hinder nutrient cycling in this moist lowland forest. [ABSTRACT FROM AUTHOR]

The relative importance of large lianas (woody vines) increased by 100% for stem enumerations conducted during the 1980s and 1990s in widely scattered Neotropical forests. We use three independent types of data to evaluate the hypothesis that lianas have increased in importance in old growth forests on Barro Colorado Island, Panama. Liana leaf lifter production and the proportion of forest-wide leaf litter composed of lianas increased between 1986 and 2002. In contrast, liana seed production and liana seedling densities were much more variable through time with particularly high levels during and immediately after El Niño years. Longer time series will be required to detect shifts in life- form composition for highly dynamic seed and seedling communities. The Barro Colorado Island leaf production data are, however, consistent with the hypothesis that lianas are increasing in importance in Neotropical forests. [ABSTRACT FROM AUTHOR]

In closed-canopy forests, plant morphology and physiology determine shade tolerance and potential growth and mortality rates; potential vital rates and ongoing gap dependence determine realized vital rates; and realized vital rates determine individual size distributions. This hypothesis was evaluated for the 73 most abundant canopy tree species from Barro Colorado Island, Panama. The percentage of recruits located in tree-fall gaps (P), sapling growth (G), and mortality (M) rates, and the coefficient of skewness of size distributions (g1) were determined from censuses of all individuals >10 mm dbh in a 50- ha plot. Seven qualitative, bivariate predictions relating g[sub1], G, M, P, and wood density (W) were evaluated. Six of the seven predictions were substantiated in pairwise analyses. A path analysis integrated all seven predictions and explained 51% of the interspecific variation in g[sub1]. Size distributions with many large individuals and a long tail of relatively rare, small individuals (g[sub1] < 0) characterized gap-dependent species with large fecundity, seed mortality, seedling mortality, G, M, and P. Size distributions with many small individuals and a long tail of relatively rare, large individuals (g[sub1] > 0) characterized shade-tolerant species with the opposite traits. The percentage of tropical tree species that require tree- fall gaps to regenerate has been estimated to range from <20% to >70% for old-growth forests. [ABSTRACT FROM AUTHOR]

Abstract: Humans hunt forest vertebrates throughout the tropics. Many preferred game species consume flowers, fruit, seeds and/or leaves, and these interactions will cause their harvest to ramify through forests. Three related issues will determine how severely the harvest of forest vertebrates influences the plant community. First, the types of species selected by hunters and the intensity of the harvest will determine which vertebrates are removed and which remain. Second, the possible presence of ecologically similar, non-game species able to expand their activities to fulfill the ecological role of heavily exploited species will determine how severely the harvest disrupts ecological relationships between the community of forest vertebrates and the community of forest plants. Finally, hunters will alter plant species composition if the harvest of vertebrates differentially affects mutualists or pests of particular plant species. Hunters will also alter plant diversity if the harvest of vertebrates disrupts ecological mechanisms that permit plant species to coexist. I examine hunter selectivity, the intensity of the hunt, possible compensation by non-game species, and the types and strengths of interactions among game species and plants for tropical forests to determine when and where these outcomes occur. [Copyright &y& Elsevier]

Examines the consequences of variable reproduction for seedling recruitment among neotropical tree species. Spatial processes affecting forest dynamics; Synchronization of trees in seed production; Temporal and spatial variation of seedling recruitment.

Examines the influence of neighboring plants on the demography of perennial shrub Ambrosia dumosa in Colorado Desert, California. Effects of neighboring plants on plant growth and survival; Documentation of differences in population growth, elasticity structure and stable size; Contributions of patch types on overall population dynamics.

Investigates the comparative phenotypic plasticity of 16 species of tropical rainforest shrubs from a Panamanian rainforest. Interactions between light treatment and the species; Similarity between shoot relative growth rates in the low light treatment; Differences between gap and understory species.