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Questions: We asked: (a) whether the strength of conspecific and heterospecific neighborhood crowding effects on focal tree survival and growth vary with neighborhood radii; and (b) if the relative strength of the effect of neighborhood interactions on tree growth and survival varies with neighborhood scale. Location: Luquillo Forest Dynamics Plot, Puerto Rico. Methods: We used tree survival and growth data and included information on species‐mean trait values related to several leaf traits, maximum height, seed mass and wood density. We incorporated a tree neighborhood modeling approach that uses an area around a focal tree with a specified radius, to describe the interactions between a focal tree and its neighbors. We constructed survival and growth models for each functional trait using a Bayesian approach, and varied the size of the radius from 5 m to 30 m, at 5‐m intervals. Results: The results suggested that the estimated effects of conspecific and heterospecific neighbors on tree performance do not vary based on the size of the neighborhood (5–30 m), suggesting that the effects of conspecific and heterospecific neighbors on the performance of a focal tree likely do not vary substantially beyond a neighborhood radius of 5 m in the Luquillo forest. In contrast, the estimated strength of the functional neighborhood (effect of neighbors based on their functional trait values) on tree performance was dependent on the neighborhood range. Our results also suggested that the effects of trait distances and trait hierarchies on tree survival and growth are acting simultaneously and at the same spatial scales. Conclusion: Findings from this study highlight the importance of spatial scale in community assembly processes, and specifically, call for increased attention when selecting the radius that defines the neighborhood around a focal tree as the selected neighborhood radius influences the community patterns discovered, and affects the conclusions about the drivers that control community assembly. [ABSTRACT FROM AUTHOR]

Abstract Forest community composition is the outcome of multiple forces, including those that increase taxonomic and functional divergence and those that promote convergence in traits. The mechanisms underlying these forces may not operate homogenously within communities; individuals of different species are never perfectly mixed, and thus, species tend to be surrounded and interact with different subsets of species. In fact, taxonomic and functional composition of neighborhoods of different focal species can be highly variable. Here, we examine whether mechanisms driving species‐level neighborhoods relate to intrinsic characteristics of focal species such as differences in life‐history and resource‐uptake strategies and in turn relate to species survival. We focus on two key characteristics: (1) seed mass, which defines a dominant axis of life‐history strategies related to stress tolerance, and (2) understory light preferences that sort species from light‐demanding pioneers to shade‐tolerant. We monitored seedling communities over 10 yr in Puerto Rico and calculated neighborhood trait dispersion in species‐level neighborhoods using seven functional traits. We examined whether species‐level characteristics, seed mass and preferred light conditions, influence patterns of functional dispersion in seedling neighborhoods using linear models. Then, we examined how species‐level functional neighborhoods impact seedling survival. We found that small‐ and large‐seeded species diverge in the type of functional neighborhoods they associate with. Large‐seeded species associate with neighbors that are more similar than expected in leaf economic traits, but more different than expected in seed mass and leaf area traits, while the opposite was found for small‐seeded species. This variation in species functional neighborhood was important in determining seedling survival. In sum, our results suggest that divergent and convergent forces do not operate homogenously over entire communities. Their relative role changes in space, and on a species‐by‐species basis, probably with a deterministic foundation linked to traits such as seed mass.

Forest community composition is the outcome of multiple forces, including those that increase taxonomic and functional divergence and those that promote convergence in traits. The mechanisms underlying these forces may not operate homogenously within communities; individuals of different species are never perfectly mixed, and thus, species tend to be surrounded and interact with different subsets of species. In fact, taxonomic and functional composition of neighborhoods of different focal species can be highly variable. Here, we examine whether mechanisms driving species-level neighborhoods relate to intrinsic characteristics of focal species such as differences in life-history and resource-uptake strategies and in turn relate to species survival. We focus on two key characteristics: (1) seed mass, which defines a dominant axis of life-history strategies related to stress tolerance, and (2) understory light preferences that sort species from light-demanding pioneers to shade-tolerant. We monitored seedling communities over 10 yr in Puerto Rico and calculated neighborhood trait dispersion in species-level neighborhoods using seven functional traits. We examined whether species-level characteristics, seed mass and preferred light conditions, influence patterns of functional dispersion in seedling neighborhoods using linear models. Then, we examined how species-level functional neighborhoods impact seedling survival. We found that small- and large-seeded species diverge in the type of functional neighborhoods they associate with. Large-seeded species associate with neighbors that are more similar than expected in leaf economic traits, but more different than expected in seed mass and leaf area traits, while the opposite was found for small-seeded species. This variation in species functional neighborhood was important in determining seedling survival. In sum, our results suggest that divergent and convergent forces do not operate homogenously over entire communities. Their relative role changes in space, and on a species-by-species basis, probably with a deterministic foundation linked to traits such as seed mass. [ABSTRACT FROM AUTHOR]

Abstract: Arbuscular mycorrhizal (AM) fungi in the soil may influence tropical tree dynamics and forest succession. The mechanisms are poorly understood, because the functional characteristics and abundances of tree species and AM fungi are likely to be codependent. We used generalized joint attribute modeling to evaluate if AM fungi are associated with three forest community metrics for a sub‐tropical montane forest in Puerto Rico. The metrics chosen to reflect changes during forest succession are the abundance of seedlings of different successional status, the amount of foliar damage on seedlings of different successional status, and community‐weighted mean functional trait values (adult specific leaf area [SLA], adult wood density, and seed mass). We used high‐throughput DNA sequencing to identify fungal operational taxonomic units (OTUs) in the soil. Model predictions showed that seedlings of mid‐ and late‐successional species had less leaf damage when the 12 most common AM fungi were abundant compared to when these fungi were absent. We also found that seedlings of mid‐successional species were predicted to be more abundant when the 12 most common AM fungi were abundant compared to when these fungi were absent. In contrast, early‐successional tree seedlings were predicted to be less abundant when the 12 most common AM fungi were abundant compared to when these fungi were absent. Finally, we showed that, among the 12 most common AM fungi, different AM fungi were correlated with functional trait characteristics of early‐ or late‐successional species. Together, these results suggest that early‐successional species might not rely as much as mid‐ and late‐successional species on AM fungi, and AM fungi might accelerate forest succession. [ABSTRACT FROM AUTHOR]

Leaf litter represents an important link between tree community composition, forest productivity and biomass, and ecosystem processes. In forests, the spatial distribution of trees and species-specific differences in leaf litter production and quality are likely to cause spatial heterogeneity in nutrient returns to the forest floor and, therefore, in the redistribution of soil nutrients. Using mapped trees and leaf litter data for 12 tree species in a subtropical forest with a well-documented history of land use, we: (1) parameterized spatially explicit models of leaf litter biomass and nutrient deposition; (2) assessed variation in leaf litter inputs across forest areas with different land use legacies; and (3) determined the degree to which the quantity and quality of leaf litter inputs and soil physical characteristics are associated with spatial heterogeneity in soil nutrient ratios (C:N and N:P). The models captured the effects of tree size and location on spatial variation in leaf litterfall (R² =3.31-0.79). For all 12 focal species, most of the leaf litter fell less than 5 m away from the source trees, generating finescale spatial heterogeneity in leaf litter inputs. Secondary forest species, which dominate areas in earlier successional stages, had lower leaf litter C:N ratios and produced less litter biomass than old-growth specialists. In contrast, P content and N:P ratios did not vary consistently among successional groups. interspecific variation in leaf litter quality translated into differences in the quantity and quality (C:N) of total leaf litter biomass inputs and among areas with different land use histories. Spatial variation in leaf litter C:N inputs was the major factor associated with heterogeneity in soil C:N ratios relative to soil physical characteristics. In contrast, spatial variation soil N:P was more strongly associated with spatial variation in topography than heterogeneity in leaf litter inputs. The modeling approach presented here can be used to generate prediction surfaces for leaf litter deposition and quality onto the forest floor, a useful tool for understanding soil-vegetation feedbacks. A better understanding of the role of leaf litter inputs from secondary vegetation in restoring soil nutrient stocks will also assist in managing expanding secondary forests in tropical regions. [ABSTRACT FROM AUTHOR]

Identifying the processes that maintain highly diverse plant communities remains a central goal in ecology. Species variation in growth and survival rates across ontogeny, represented by tree size classes and life history stage-specific niche partitioning, are potentially important mechanisms for promoting forest diversity. However, the role of ontogeny in mediating competitive dynamics and promoting functional diversity is not well understood, particular in high-diversity systems such as tropical forests. The interaction between interspecific functional trait variation and ontogenetic shifts in competitive dynamics may yield insights into the ecophysiological mechanisms promoting community diversity. We investigated how functional trait (seed size, maximum height, SLA, leaf N, and wood density) associations with growth, survival, and response to competing neighbors differ among seedlings and two size classes of trees in a subtropical rain forest in Puerto Rico. We used a hierarchical Bayes model of diameter growth and survival to infer trait relationships with ontogenetic change in competitive dynamics. Traits were more strongly associated with average growth and survival than with neighborhood interactions, and were highly consistent across ontogeny for most traits. The associations between trait values and tree responses to crowding by neighbors showed significant shifts as trees grew. Large trees exhibited greater growth as the difference in species trait values among neighbors increased, suggesting traitassociated niche partitioning was important for the largest size class. Our results identify potential axes of niche partitioning and performance-equalizing functional trade-offs across ontogeny, promoting species coexistence in this diverse forest community. [ABSTRACT FROM AUTHOR]

The article presents the study on the relation of human and natural disturbances to forest regeneration and recovery in Puerto Rico. Researchers monitored the seedlings over ten years and found that the hurricane disturbance has a great impact on seed dynamics and seedling composition. However, it was observed that hurricane helps preserve the signature of land use history and promotes the growth and survival of seedlings of secondary-forest species.

1. Many forests experience periodic, large-scale disturbances, such as hurricanes and cyclones, which open the forest canopy, causing dramatic changes in understorey light conditions and seedling densities. Thus, in hurricane-impacted forests, large variations in abiotic and biotic conditions likely shape seedling dynamics, which in turn will contribute to patterns of forest recovery. 2. We monitored 13 836 seedlings of 82 tree and shrub species over 10 years following Hurricane Georges in 1998 in a subtropical, montane forest in Puerto Rico. We quantified changes in the biotic and abiotic environment of the understorey and linked seedling dynamics to changes in canopy openness and seedling density, and to spatial variation in soil type, topography and tree density. 3. Canopy openness was highest when first measured after Hurricane Georges and dropped significantly within c. 3 years, while seedling densities remained high for c. 5 years post-hurricane. When all species and census intervals were analysed together, generalized linear mixed effects models revealed that canopy openness, seedling and adult tree densities were significant drivers of seedling survival. 4. The relative importance of abiotic and biotic factors changed over time. Separate analyses for each census interval revealed that canopy openness was a significant predictor of survival only for the first census interval, with lower survival at the highest levels of canopy openness. The effect of conspecific seedling density was significant in all intervals except the first, and soil type only in the final census interval. 5. When grouping species into life-history guilds based on adult tree susceptibility to hurricane damage, we found clear differences among guilds in the effects of biotic and abiotic factors on seedling survival. Seedlings of hurricane-susceptible and intermediate guilds were more strongly influenced by canopy openness, while seedlings of the hurricane-resistant group were less affected by conspecific seedling density. Individual species-level analyses for 12 common species, however, showed considerable variation among species within guilds. 6. Synthesis. Our results suggest that hurricanes shape species composition by altering understorey conditions that differentially influence the success of seedlings. Thus, predicted increases in the intensity and frequency of hurricanes in the Caribbean will likely alter seedling dynamics and ultimately the species composition in hurricane-impacted forests. [ABSTRACT FROM AUTHOR]

Forests are often subject to multiple, compounded disturbances, representing both natural and human-induced processes. Predicting forest dynamics requires that we consider how these disturbances interact to affect species demography. Here we present results of an individual-based, spatially explicit forest simulator that we developed to analyze the compounded effects of hurricane disturbance and land use legacies on the dynamics of a subtropical forest. We used data from the 16-ha Luquillo Forest Dynamics Plot in Puerto Rico, together with a reconstruction of historical wind damage, to parameterize the simulator. We used the model to ask two questions. (1) What are the implications of variation in hurricane frequency and severity for the long-term dynamics of forest composition, diversity, and structure? Both storm severity and frequency had striking effects on forest dynamics, composition, and structure. The periodicity of disturbance also played an important role, with periods of high hurricane activity fostering the establishment of species that may become rare in the absence of severe storms and quiescent periods allowing these species to reach reproductive size. Species responses to hurricane disturbance could not be predicted from their life history attributes. However, species perceived to be primary forest species exhibited low temporal variation in abundance through the simulations. (2) How do hurricanes and legacies from human land use interact to determine community structure and composition? Our results suggest that, over time, regardless of the storm regime, land use legacies will become less apparent but will lead to a forest community that contains a mixture of secondary and primary forest species formerly dominant in areas of different land use. In the long term, hurricane disturbance generated two communities with slightly greater similarity than those not subject. to storms. Thus, the inclusion of hurricane disturbance does not alter the prediction that land use legacies in this tropical forest will diminish over time. Our simulations also highlight the contingent effects of human legacies on subsequent community dynamics, including the response to hurricane disturbance, therefore supporting the notion that compounded disturbances can interact in ways that cannot be predicted by the study of single disturbances. The widespread importance of land use as a large-scale disturbance makes it imperative that it be addressed as a fundamental ecological process. [ABSTRACT FROM AUTHOR]

We present a likelihood-based regression method that was developed to analyze the effects of neighborhood competitive interactions and hurricane damage on tree growth and survival. The purpose of the method is to provide robust parameter estimates for a spatially explicit forest simulator and to gain insight into the processes that drive the patterns of species abundance in tropical forests. We test the method using census data from the 16-ha Luquillo Forest Dynamics Plot in Puerto Rico and describe effects of the spatial configuration, sizes, and species of neighboring trees on the growth and survival of 12 dominant tree species representing a variety of life history strategies. Variation in size-dependent growth and mortality suggests a complex relationship between size, growth, and survival under different regimes of light availability. Crowding effects on growth and survival appear to be idiosyncratic to each individual species, and with the exception of pioneers, there is little commonality among species that share similar life histories. We also explain the implications of differential susceptibility to hurricane damage on species' growth and survival and on their ability to respond to damage to neighboring trees. Tree species in the Luquillo Forest Dynamics Plot differ strikingly in both their susceptibility to hurricane disturbance and the nature of their recovery from wind disturbance, through response of both adult plants and juveniles to enhanced resource availability. At the stand level, intense competitive thinning of densely packed saplings that grew after hurricane damage accounted for the majority of post-hurricane mortality, particularly for shade-intolerant species. At the individual species level, effects of previous hurricane damage on growth and survival depended primarily on variation in the quantity and quality of hurricane damage sustained "by target species and their interaction with life history characteristics of these individual species. Finally, we compare models that make different assumptions about the effects of competing species on tree growth and survival (e.g., equivalence of competitors vs. distinct species-specific effects). Size effects alone could not account for growth and survival for the majority of target species. Our results also demonstrate that competing species have distinct per capita effects on growth of dominant target species. In contrast, we found moderate support for a model that assumed functional equivalence of competitors on survival. [ABSTRACT FROM AUTHOR]