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Background and Aims Understanding shifts in the demographic and functional composition of forests after major natural disturbances has become increasingly relevant given the accelerating rates of climate change and elevated frequency of natural disturbances. Although plant demographic strategies are often described across a slow–fast continuum, severe and frequent disturbance events influencing demographic processes may alter the demographic trade-offs and the functional composition of forests. We examined demographic trade-offs and the shifts in functional traits in a hurricane-disturbed forest using long-term data from the Luquillo Forest Dynamics Plot (LFPD) in Puerto Rico. Methods We analysed information on growth, survival, seed rain and seedling recruitment for 30 woody species in the LFDP. In addition, we compiled data on leaf, seed and wood functional traits that capture the main ecological strategies for plants. We used this information to identify the main axes of demographic variation for this forest community and evaluate shifts in community-weighted means for traits from 2000 to 2016. Key Results The previously identified growth–survival trade-off was not observed. Instead, we identified a fecundity–growth trade-off and an axis representing seedling-to-adult survival. Both axes formed dimensions independent of resprouting ability. Also, changes in tree species composition during the post-hurricane period reflected a directional shift from seedling and tree communities dominated by acquisitive towards conservative leaf economics traits and large seed mass. Wood specific gravity, however, did not show significant directional changes over time. Conclusions Our study demonstrates that tree demographic strategies coping with frequent storms and hurricane disturbances deviate from strategies typically observed in undisturbed forests, yet the shifts in functional composition still conform to the expected changes from acquisitive to conservative resource-uptake strategies expected over succession. In the face of increased rates of natural and anthropogenic disturbance in tropical regions, our results anticipate shifts in species demographic trade-offs and different functional dimensions. [ABSTRACT FROM AUTHOR]

Severe droughts have led to lower plant growth and high mortality in many ecosystems worldwide, including tropical forests. Drought vulnerability differs among species, but there is limited consensus on the nature and degree of this variation in tropical forest communities. Understanding species‐level vulnerability to drought requires examination of hydraulic traits since these reflect the different strategies species employ for surviving drought.Here, we examined hydraulic traits and growth reductions during a severe drought for 12 common woody species in a wet tropical forest community in Puerto Rico to ask: Q1. To what extent can hydraulic traits predict growth declines during drought? We expected that species with more hydraulically vulnerable xylem and narrower safety margins (SMP50) would grow less during drought. Q2. How does species successional association relate to the levels of vulnerability to drought and hydraulic strategies? We predicted that early‐ and mid‐successional species would exhibit more acquisitive strategies, making them more susceptible to drought than shade‐tolerant species. Q3. What are the different hydraulic strategies employed by species and are there trade‐offs between drought avoidance and drought tolerance? We anticipated that species with greater water storage capacity would have leaves that lose turgor at higher xylem water potential and be less resistant to embolism forming in their xylem (P50).We found a large range of variation in hydraulic traits across species; however, they did not closely capture the magnitude of growth declines during drought. Among larger trees (≥10 cm diameter at breast height—DBH), some tree species with high xylem embolism vulnerability (P50) and risk of hydraulic failure (SMP50) experienced substantial growth declines during drought, but this pattern was not consistent across species. We found a trade‐off among species between drought avoidance (capacitance) and drought tolerating (P50) in this tropical forest community. Hydraulic strategies did not align with successional associations. Instead, some of the more drought‐vulnerable species were shade‐tolerant dominants in the community, suggesting that a drying climate could lead to shifts in long‐term forest composition and function in Puerto Rico and the Caribbean. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Summary: Rapid changes in climate and disturbance regimes, including droughts and hurricanes, are likely to influence tropical forests, but our understanding of the compound effects of disturbances on forest ecosystems is extremely limited. Filling this knowledge gap is necessary to elucidate the future of these ecosystems under a changing climate.We examined the relationship between hurricane response (damage, mortality, and resilience) and four hydraulic traits of 13 dominant woody species in a wet tropical forest subject to periodic hurricanes.Species with high resistance to embolisms (low P50 values) and higher safety margins (SMP50) were more resistant to immediate hurricane mortality and breakage, whereas species with higher hurricane resilience (rapid post‐hurricane growth) had high capacitance and P50 values and low SMP50. During 26 yr of post‐hurricane recovery, we found a decrease in community‐weighted mean values for traits associated with greater drought resistance (leaf turgor loss point, P50, SMP50) and an increase in capacitance, which has been linked with lower drought resistance.Hurricane damage favors slow‐growing, drought‐tolerant species, whereas post‐hurricane high resource conditions favor acquisitive, fast‐growing but drought‐vulnerable species, increasing forest productivity at the expense of drought tolerance and leading to higher overall forest vulnerability to drought. [ABSTRACT FROM AUTHOR]

The distribution of Bornean heath forest on white sand soils is believed to be due to element limitation and soil acidity.To determine the impact of both element limitation and soil acidity on tropical heath forest, we established a soil fertilization experiment to investigate the impact that increased soil N availability and reduced soil pH (using lime/CaCO3) had on a range of elements in tree leaves. We hypothesized that alterations in soil resource availability would cause changes in the tree leaf N:P ratio and concentrations of other elements and these changes would be influenced by species' functional strategies.The experiment was carried out in a Bornean heath forest on infertile soil over a 2‐year period. We selected 10 common tree species, spanning acquisitive to conservative strategies and tested whether tree species functional strategies influenced tree leaf elemental concentrations after the fertilization.Leaf N:P ratios showed considerable differences among co‐occurring species. Overall, we found that soil N addition treatments increased leaf N concentration, although leaf N:P ratios were not affected. Changes in leaf Al, Fe and S concentrations were correlated with species functional strategy: conservative species showed a greater increase in leaf Fe compared with acquisitive species, whereas acquisitive species showed a greater increase in leaf Al but a decrease in leaf S compared with conservative species.Synthesis. We show that soil elements uptake differs between acquisitive and conservative species and that acquisitive species may not take up soil elemental resources more effectively than conservative species. We suggest that the greater Fe acquisition by conservative species, in comparison to acquisitive species, might be due to a stress tolerance strategy. The overall increase in leaf N showed that, in this nutrient‐poor forest, N is a fundamental requirement irrespective of species functional strategies. Given the increased leaf Al and Fe concentration after liming, we also suggest that these elements are important, and possibly limiting (Fe), in this heath forest. [ABSTRACT FROM AUTHOR]

The study of community spatial structure is central to understanding diversity patterns over space and species co‐occurrence at local scales. Although most analytical approaches consider horizontal and vertical dimensions separately, in this study we introduce a three‐dimensional spatial analysis that simultaneously includes horizontal and vertical species associations. Using tree census data (2000–2016) and allometries from the Luquillo forest plot in Puerto Rico, we show that spatial organization becomes less random over time as the forest recovered from land‐use legacy effects and hurricane disturbance. Tree species vertical segregation is predominant in the forest with almost all species that co‐occur in the horizontal plane avoiding each other in the vertical dimension. Horizontal segregation is less common than vertical, whereas three‐dimensional aggregation (a proxy for direct tree competition) is the least frequent type of spatial association. Furthermore, dominant species are involved in more non‐random spatial associations, implying that species co‐occurrence is facilitated by species segregation in space. This novel three‐dimensional analysis allowed us to identify and quantify tree species spatial distributions, how interspecific competition was reduced through forest structure, and how it changed over time after disturbance, in ways not detectable from two‐dimensional analyses alone. [ABSTRACT FROM AUTHOR]

Legumes provide an essential service to ecosystems by capturing nitrogen from the atmosphere and delivering it to the soil, where it may then be available to other plants. However, this facilitation by legumes has not been widely studied in global tropical forests. Demographic data from 11 large forest plots (16–60 ha) ranging from 5.25° S to 29.25° N latitude show that within forests, leguminous trees have a larger effect on neighbor diversity than non-legumes. Where soil nitrogen is high, most legume species have higher neighbor diversity than non-legumes. Where soil nitrogen is low, most legumes have lower neighbor diversity than non-legumes. No facilitation effect on neighbor basal area was observed in either high or low soil N conditions. The legume–soil nitrogen positive feedback that promotes tree diversity has both theoretical implications for understanding species coexistence in diverse forests, and practical implications for the utilization of legumes in forest restoration. Xu et al. examine the effect of leguminous trees on neighbor diversity across 11 plots in tropical forests around the world, and find that in high soil nitrogen conditions, most legume species have higher neighbor diversity than non-legumes, and vice versa where soil nitrogen is low. Their results have practical implications for the utilization of legumes in forest restoration. [ABSTRACT FROM AUTHOR]

Projected increases in cyclonic storm intensity under a warming climate will have profound effects on forests, potentially changing these ecosystems from carbon sinks to sources. Forecasting storm impacts on these ecosystems requires consideration of risk factors associated with storm meteorology, landscape structure, and forest attributes. Here we evaluate risk factors associated with damage severity caused by Hurricanes María and Irma across Puerto Rican forests. Using field and remote sensing data, total forest aboveground biomass (AGB) lost to the storms was estimated at 10.44 (±2.33) Tg, ca. 23% of island-wide pre-hurricane forest AGB. Storm-related rainfall was a stronger predictor of forest damage than maximum wind speeds. Soil water storage capacity was also an important risk factor, corroborating the influence of rainfall on forest damage. Expected increases of 20% in hurricane-associated rainfall in the North Atlantic highlight the need to consider how such shifts, together with high speed winds, will affect terrestrial ecosystems. [ABSTRACT FROM AUTHOR]

Background and aims: Whilst several studies have shown that edaphic variability influences species composition in nutrient-poor tropical forests, the determinants of local species distributions and, in particular, how these change from younger to mature individuals in such forests are still under debate, and have been poorly explored in tropical heath forests that are among the least fertile tropical forest ecosystems. Methods: We investigated the influence of soil fertility and topography on a Bornean heath forest species composition, α-, β-diversity and tree size structure among size classes by recording all trees ≥1 cm DBH in 16 forest plots totalling 0.36 ha. Results: Tree species distributions generally followed gradients in available Al and soil depth; α- and β-diversity were linked to soil depth, and to some extent also to pH and the H:Al ratio. In contrast, forest structural attributes (basal area and stem density) were negatively correlated with both available and total P and a wider suite of soil nutrients, although trees ≥10 cm DBH were positively correlated with total P. Conclusion: Our study shows that heath forest species distribution, richness and structure is related to both edaphic and topographic characteristics and that soil acidity might have a strong influence in shaping these forests' features. Among size classes, small trees are less influenced by soil and topography, whereas the sensitivity to these variables increases with tree size. We thus highlight that multiple edaphic factors influence different aspects of tropical forest structure, including different tree life stages, and species composition. [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]

Tropical forests play a critical role in carbon and water cycles at a global scale. Rapid climate change is anticipated in tropical regions over the coming decades and, under a warmer and drier climate, tropical forests are likely to be net sources of carbon rather than sinks. However, our understanding of tropical forest response and feedback to climate change is very limited. Efforts to model climate change impacts on carbon fluxes in tropical forests have not reached a consensus. Here, we use the Ecosystem Demography model (ED2) to predict carbon fluxes of a Puerto Rican tropical forest under realistic climate change scenarios. We parameterized ED2 with species-specific tree physiological data using the Predictive Ecosystem Analyzer workflow and projected the fate of this ecosystem under five future climate scenarios. The model successfully captured interannual variability in the dynamics of this tropical forest. Model predictions closely followed observed values across a wide range of metrics including aboveground biomass, tree diameter growth, tree size class distributions, and leaf area index. Under a future warming and drying climate scenario, the model predicted reductions in carbon storage and tree growth, together with large shifts in forest community composition and structure. Such rapid changes in climate led the forest to transition from a sink to a source of carbon. Growth respiration and root allocation parameters were responsible for the highest fraction of predictive uncertainty in modeled biomass, highlighting the need to target these processes in future data collection. Our study is the first effort to rely on Bayesian model calibration and synthesis to elucidate the key physiological parameters that drive uncertainty in tropical forests responses to climatic change. We propose a new path forward for model-data synthesis that can substantially reduce uncertainty in our ability to model tropical forest responses to future climate. [ABSTRACT FROM AUTHOR]

Impacts of climate change require that society urgently develops ways to reduce amounts of carbon in the atmosphere. Tropical forests present an important opportunity, as they take up and store large amounts of carbon. It is often suggested that forests with high biodiversity have large stocks and high rates of carbon uptake. Evidence is, however, scattered across geographic areas and scales, and it remains unclear whether biodiversity is just a co-benefit or also a requirement for the maintenance of carbon stocks and uptake. Here, we perform a quantitative review of empirical studies that analyzed the relationships between plant biodiversity attributes and carbon stocks and carbon uptake in tropical forests. Our results show that biodiversity attributes related to species, traits or structure significantly affect carbon stocks or uptake in 64% of the evaluated relationships. Average vegetation attributes (community-mean traits and structural attributes) are more important for carbon stocks, whereas variability in vegetation attributes ( i.e., taxonomic diversity) is important for both carbon stocks and uptake. Thus, different attributes of biodiversity have complementary effects on carbon stocks and uptake. These biodiversity effects tend to be more often significant in mature forests at broad spatial scales than in disturbed forests at local spatial scales. Biodiversity effects are also more often significant when confounding variables are not included in the analyses, highlighting the importance of performing a comprehensive analysis that adequately accounts for environmental drivers. In summary, biodiversity is not only a co-benefit, but also a requirement for short- and long-term maintenance of carbon stocks and enhancement of uptake. Climate change policies should therefore include the maintenance of multiple attributes of biodiversity as an essential requirement to achieve long-term climate change mitigation goals. [ABSTRACT FROM AUTHOR]

Aim Determining the drivers of species rarity is fundamental for understanding and conserving biodiversity. Rarity of a given species within its community may arise due to exclusion by other ecologically similar species. Conversely, rare species may occupy habitats that are rare in the landscape or they may be ill-suited to all available habitats. The first mechanism would lead to common and rare species occupying similar ecological space defined by functional traits. The second mechanism would result in common and rare species occupying dissimilar ecological space and spatial aggregation of rare species, either because they are specialists in rare habitats or because rare species tend to be dispersal limited. Here, we quantified the contribution of locally rare species to community functional richness and the spatial aggregation of species across tree communities world-wide to address these hypotheses. Location Asia and the Americas. Time period 2002 to 2012 (period that considers the censuses for the plots used). Major taxa studied Angiosperm and Gymnosperm trees. Methods We compiled a dataset of functional traits from all the species present in eight tree plots around the world to evaluate the contribution of locally rare species to tree community functional richness using multi- and univariate approaches. We also quantified the spatial aggregation of individuals within species at several spatial scales as it relates to abundance. Results Locally rare tree species in temperate and tropical forests tended to be functionally unique and are consistently spatially clustered. Furthermore, there is no evidence that this pattern is driven by pioneer species being locally rare. Main conclusions This evidence shows that locally rare tree species disproportionately contribute to community functional richness, and we can therefore reject the hypothesis that locally rare species are suppressed by ecologically similar, but numerically dominant, species. Rather, locally rare species are likely to be specialists on spatially rare habitats or they may be ill-suited to the locally available environments. [ABSTRACT FROM AUTHOR]

Our understanding of the long-lasting effects of human land use on soil fungal communities in tropical forests is limited. Yet, over 70% of all remaining tropical forests are growing in former agricultural or logged areas. We investigated the relationship among land use history, biotic and abiotic factors, and soil fungal community composition and diversity in a second-growth tropical forest in Puerto Rico. We coupled high-throughput DNA sequencing with tree community and environmental data to determine whether land use history had an effect on soil fungal community descriptors. We also investigated the biotic and abiotic factors that underlie such differences and asked whether the relative importance of biotic (tree diversity, basal tree area, and litterfall biomass) and abiotic (soil type, pH, iron, and total carbon, water flow, and canopy openness) factors in structuring soil fungal communities differed according to land use history. We demonstrated long-lasting effects of land use history on soil fungal communities. At our research site, most of the explained variation in soil fungal composition ( R2 = 18.6%), richness ( R2 = 11.4%), and evenness ( R2 = 10%) was associated with edaphic factors. Areas previously subject to both logging and farming had a soil fungal community with lower beta diversity and greater evenness of fungal operational taxonomic units (OTUs) than areas subject to light logging. Yet, fungal richness was similar between the two areas of historical land use. Together, these results suggest that fungal communities in disturbed areas are more homogeneous and diverse than in areas subject to light logging. Edaphic factors were the most strongly correlated with soil fungal composition, especially in areas subject to light logging, where soils are more heterogenous. High functional tree diversity in areas subject to both logging and farming led to stronger correlations between biotic factors and fungal composition than in areas subject to light logging. In contrast, fungal richness and evenness were more strongly correlated with biotic factors in areas of light logging, suggesting that these metrics might reflect long-term associations in old-growth forests. The large amount of unexplained variance in fungal composition suggests that these communities are structured by both stochastic and niche assemblage processes. [ABSTRACT FROM AUTHOR]

1. Tropical forest tree diversity has been hypothesized to be maintained via the attraction of density responsive and species-specific enemies. Tests of this hypothesis usually assume a linear relationship between enemy pressure (amount of damage and enemy richness) and seedling or tree density. However, enemy pressure is likely to change nonlinearly with local seedling abundance and community scale tree abundance if enemies are characterized by nonlinear functional responses. 2. We examined the abiotic and biotic factors associated with richness of above-ground enemies and foliar damage found in tree seedlings in a tropical forest in Puerto Rico. Rather than identify specific enemies targeting these seedlings, we used damage morphotypes, a paleo-ecological method, to derive a proxy for enemy species richness. 3. We found that the relationships between local and (conspecific seedling density) community scale (conspecific basal area of adult trees) abundance and both richness of above-ground enemies and foliar damage were hump-shaped. Seedlings of tree species existing at intermediate levels of abundance, at both local and community scales, suffered more damage and experienced pressure from a greater diversity of enemies than those existing at high or low densities. 4. We hypothesized that greater damage at intermediate abundance level could arise from a rich mixture of generalist and specialist enemies targeting seedlings of intermediate abundance tree species. Consistent with this hypothesis, we found that generalist enemies were more diverse on species at rare or intermediate abundance relative to common tree species. However, specialist enemies showed no significant trend across tree species abundance at either the local or community scales. 5. Synthesis. Our results suggest that interspecific variation in tree species abundance leads to differences in the magnitude and type of damage tropical tree seedlings suffer. This variation leads to a nonlinear, hump-shaped relationship between species abundance and enemy damage, highlighting fruitful directions for further development of species coexistence theory. [ABSTRACT FROM AUTHOR]

• Experimental increase of soil pH in tropical heath forest increase mesofaunal activity. • Modification of soil pH and N availability had weak effect on organic matter decomposition. • Organic matter decomposition rate in tropical heath forest calculated with standard tea bags is similar to temperate forest. Decomposition is a key process for rain forest nutrient cycling, and this may be altered by the increasing rate of deposition of reactive nitrogen on rain forests. Tropical heath forests are characterised by slow litter decomposition rates due to low quality litter, along with acidic soil pH and low availability of soil nitrogen (N). To investigate whether soil N or pH is most important in regulating tropical heath forest organic matter decomposition, we used a factorial N and CaCO 3 addition experiment, and measured wood and leaf litter decomposition rates and mesofaunal activity (using bait-lamina sticks and numbers of worm casts). Our heath forest site in the Kabili-Sepilok Forest Reserve (Sabah, Malaysia) had a decomposition rate (calculated using standard tea material) comparable to that of temperate forest soils. Overall, the experimental modification of both soil pH and available N had weak effects on decomposition rates, although different materials responded differently to the experimental treatments. There was a clear, but transient, increase in number of worm casts in the CaCO 3 and N + CaCO 3 treatment. Although we obtained some evidence that low soil acidity reduced decomposer activity, it may take longer than one year for the decomposer community, and associated processes, to be influenced by the experimentally altered edaphic conditions. [ABSTRACT FROM AUTHOR]

Species composition and community structure in Neotropical forests have been severely affected by increases in climate change and disturbance. Among the most conspicuous changes is the proliferation of lianas. These increases have affected not only the carbon storage capacity of forests but also tree dynamics by reducing tree growth and increasing mortality. Despite the importance of lianas in Neotropical forests, most of the studies on lianas have focused on adult stages, ignoring dynamics at the seedlings stage. Here, we asked whether observed increases in liana abundance are associated with a demographic advantage that emerges early in liana ontogeny and with decreased precipitation and increased disturbance. To test this, we compared patterns of growth and survival between liana seedlings and tree seedlings using a long‐term data set of seedling plots from a subtropical wet forest in Puerto Rico, USA. Then, we examined the effect of precipitation and land use history on these demographic variables. We found evidence for liana seedling survival advantage over trees, but no growth advantages. This survival advantage exhibited significant temporal variation linked with patterns of rainfall, as well as differences associated with land‐use history in the study area. Furthermore, we found that neighborhood density has a negative effect on liana survival and growth. Our results indicate that liana proliferation is likely related to a survival advantage that emerges in early stages and is influenced by climatic conditions and past disturbance. Predicted climatic changes in rainfall patterns, including more frequent and severe droughts, together with increases in disturbance, could have a significant effect on seedling tropical communities by favoring lianas. [ABSTRACT FROM AUTHOR]

As cyclonic wind storms (hurricanes and typhoons) increase in frequency and intensity with climate change, it is important to understand their effects on the populations and communities of tropical trees they impact. Using tree demographic data from four large, tropical forest dynamics plots that differ in cyclonic storm frequency, we compare tree population and community dynamics. Additionally, we assess the effect of cyclonic storms on three functional traits, specific leaf area, wood density, and tree height of the dynamic tree assemblages. Mortality, growth and recruitment rates and the intrinsic rates of population growth of species differed across the plots, and were most dynamic, especially for stems 1–2 cm in diameter, at the plot which had an intermediate level of cyclonic storm frequency. Functional assemblages of species had the greatest degree of temporal variation in relation to disturbance, as measured by the change in functional divergence for the two plots with more intermediate cyclonic storm recurrence. Therefore, cyclonic storms affecting these plots generally have a greater effect on forest composition and dynamism than comparable cyclonic storms do on the plot which experiences cyclonic storms more frequently. Thus, we provide some evidence that community-wide demographic resistance to cyclonic storms is generally lower at an intermediate frequency of storms. While cyclonic storm strength and timing are important determinants of the within forest variation in tree dynamics and functional trait assemblages, we also show that cyclonic storm timing and frequency shapes tropical forest dynamics and functional composition across forests. We conclude that, over a given time interval, sites with intermediate levels of damaging cyclonic wind disturbance express a greater potential for life-history variation in the forest community, when compared to sites with less or more frequent disturbance. [ABSTRACT FROM AUTHOR]