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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]

Over the past two decades, liana density and basal area have been increasing in many tropical forests, which has profound consequences for forest diversity and functioning. One hypothesis to explain increasing lianas is elevated nutrient deposition in tropical forests resulting from fossil fuels, agricultural fertilizer, and biomass burning. We tested this hypothesis by surveying all lianas ≥1 cm in diameter (n = 3,967) in 32 plots in a fully factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment in a mature tropical forest in central Panama. We conducted the nutrient‐addition experiment from 1998 until present and we first censused lianas in 2013 and then again in 2018. After 20 yr of nutrient addition (1998–2018), liana density, basal area, and rarefied species richness did not differ significantly among any of the nutrient‐addition and control treatments. Moreover, nutrient addition in the most recent 5 yr of the experiment did not affect liana relative growth, recruitment, or mortality rates. From 2013 until 2018, liana density, basal area, and species richness increased annually by 1.6%, 1.4%, and 2.4%, respectively. Nutrient addition did not influence these increases. Our findings indicate that nutrient deposition does not explain increasing lianas in this tropical forest. Instead, increases in tree mortality and disturbance, atmospheric carbon dioxide, drought frequency and severity, and hunting pressure may be more likely explanations for the increase in lianas in tropical forests. [ABSTRACT FROM AUTHOR]

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

Abstract: Abiotic constraints and biotic interactions act simultaneously to shape communities. However, these community assembly mechanisms are often studied independently, which can limit understanding of how they interact to affect species dynamics and distributions. We develop a hierarchical Bayesian neighborhood modeling approach to quantify the simultaneous effects of topography and crowding by neighbors on the growth of 124,704 individual stems ≥1 cm DBH for 1,047 tropical tree species in a 25‐ha mapped rainforest plot in Amazonian Ecuador. We build multi‐level regression models to evaluate how four key functional traits (specific leaf area, maximum tree size, wood specific gravity and seed mass) mediate tree growth response to topography and neighborhood crowding. Tree growth is faster in valleys than on ridges and is reduced by neighborhood crowding. Topography and crowding interact to influence tree growth in ~10% of the species. Specific leaf area, maximum tree size and seed mass are associated with growth responses to topography, but not with responses to neighborhood crowding or with the interaction between topography and crowding. In sum, our study reveals that topography and neighborhood crowding each influence tree growth in tropical forests, but act largely independently in shaping species distributions. While traits were associated with species response to topography, their role in species response to neighborhood crowding was less clear, which suggests that trait effects on neighborhood dynamics may depend on the direction (negative/positive) and degree of symmetry of biotic interactions. Our study emphasizes the importance of simultaneously assessing the individual and interactive role of multiple mechanisms in shaping species dynamics in high diversity tropical systems. [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]

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]

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]

Patterns of seed dispersal and seed mortality influence the spatial structure of plant communities and the local coexistence of competing species. Most seeds are dispersed in proximity to the parent tree, where mortality is also expected to be the highest, because of competition with siblings or the attraction of natural enemies. Whereas distance‐dependent mortality in the seed‐to‐seedling transition was often observed in tropical forests, few studies have attempted to estimate the shape of the survival‐distance curves, which determines whether the peak of seedling establishment occurs away from the parent tree (Janzen–Connell pattern) or if the peak attenuates but remains at the parent location (Hubbell pattern). In this study, we inferred the probability density of seed dispersal and two stages of seedling establishment (new recruits, and seedlings 20 cm or taller) with distance for 24 tree species present in the 50‐ha Forest Dynamics Plot of Barro Colorado Island, Panama. Using data from seed traps, seedling survey quadrats, and tree‐census records spanning the 1988–2014 period, we fit hierarchical Bayesian models including parameters for tree fecundity, the shape of the dispersal kernel, and overdispersion of seed or seedling counts. We combined predictions from multiple dispersal kernels to obtain more robust inferences. We find that Hubbell patterns are the most common and Janzen–Connell patterns are very rare among those species; that distance‐dependent mortality may be stronger in the seed stage, in the early recruit stage, or comparable in both; and that species with larger seeds experience less overall mortality and less distance‐dependent mortality. Finally, we describe how this modeling approach could be extended at a community scale to include less abundant species. [ABSTRACT FROM AUTHOR]