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Canopy gaps are widely recognized as being crucial for maintaining the diversity of forest tree communities. But empirical studies have found mixed results because the differences in diversity between individual gaps and non-gaps are often small and statistically undetectable. One overlooked factor, however, is how small individual gap versus non-gap differences may accumulate across sites and potentially have a large effect on forest diversity at the plot scale. Our study investigated sapling richness, density, and composition in 124 treefall gaps, and 200 non-gap sites in the 50-ha tropical forest plot at Barro Colorado Island (BCI), Panama. Additionally, we analyzed species accumulation curves to understand how species richness increases with increasing stem numbers. We observed that sapling richness and density were only slightly higher in gaps 7 years after formation and statistically indistinguishable from non-gaps after 12 years. However, species accumulation curves across multiple gaps were substantially higher than those across non-gaps. Species composition showed small differences between individual gaps and non-gaps but differed significantly between collections of gaps and non-gaps. Specifically, 55 species specialized in 7-year-old gaps compared with 24 in non-gaps; of these, 23 gap-specialized species and zero non-gap species were pioneers. Our results indicate that tree species richness is higher in gaps because of both higher stem density and the presence of gap-specialized species. Our study has finally provided compelling evidence to support the idea that gaps enhance the overall diversity of tropical forest tree communities., (© 2024 The Author(s). Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Litter layer, serving as the "skin" of forest soil, plays a crucial role in conserving water resources and maintaining soil and water conservation. We analyzed the relationship of tree species richness, community weighted mean traits, and functional diversity with the standing mass, maximum water holding rate, and effective water sto-rage capacity of litters from various tree species including Liquidambar formosana , Mytilaria laosensis , Castanopsis sclerophylla , Castanopsis hystrix , Cunninghamia lanceolata , Pinus massoniana , Fokienia hodginsii , Taxus wallichiana and their combinations of mixed forests in subtropical region. The results showed that across various tree species combinations, the ranges of maximum water holding rate, standing litter mass and effective water storage capacity of undecomposed layer were 0-419%, 0-0.58 t·hm -2 , and 0-1.66 t·hm -2 , respectively. For the semi-decomposition layer, these values spanned in 0-375%, 0-6.14 t·hm -2 , and 0-16.03 t·hm -2 , respectively. Tree species richness and community weighted mean specific leaf area had significantly positive effects on standing mass of litter and effective water storage capacity, while community weighted mean leaf N content had significantly negative effect on standing mass of litter. The maximum water holding rate increased with the increases of functional diversity of specific leaf area and community weighted mean specific leaf area, decreased with the increase of community weighted mean leaf thickness. Results of structural equation model showed that tree species richness increased litter water holding capacity by increasing functional diversity of specific leaf area. The community weighted mean specific leaf area increased the water holding capacity of litter layer by increasing standing mass of litter and the maximum water holding rate. It is necessary to consider planting mixed forest with higher community weighted mean specific leaf area in the management of subtropical artificial forest, so as to improve the water holding capacity of litter layer.

Logged and disturbed forests are often viewed as degraded and depauperate environments compared with primary forest. However, they are dynamic ecosystems 1 that provide refugia for large amounts of biodiversity 2,3 , so we cannot afford to underestimate their conservation value 4 . Here we present empirically defined thresholds for categorizing the conservation value of logged forests, using one of the most comprehensive assessments of taxon responses to habitat degradation in any tropical forest environment. We analysed the impact of logging intensity on the individual occurrence patterns of 1,681 taxa belonging to 86 taxonomic orders and 126 functional groups in Sabah, Malaysia. Our results demonstrate the existence of two conservation-relevant thresholds. First, lightly logged forests (<29% biomass removal) retain high conservation value and a largely intact functional composition, and are therefore likely to recover their pre-logging values if allowed to undergo natural regeneration. Second, the most extreme impacts occur in heavily degraded forests with more than two-thirds (>68%) of their biomass removed, and these are likely to require more expensive measures to recover their biodiversity value. Overall, our data confirm that primary forests are irreplaceable 5 , but they also reinforce the message that logged forests retain considerable conservation value that should not be overlooked., (© 2024. The Author(s).)

Here we studied the entire Atlantic Forest hotspot to investigate whether the effect of different environmental predictors depends on the phylogenetic extension and the biogeographical history of different Atlantic Forest sectors. We used occurrence data of 3,183 plant species with arboreal or arborescent habits. We reconstructed climatic stability across 120,000 years using the Random Forest method. Then, we compared the effect of biogeographical history, topographic, and climatic variables on species richness and phylogenetic diversity using Geographically Weighted Regression (GWR) models. Niche conservatism drives the strength and direction of environmental correlates with tree diversity, interacting with the biogeographical and phylogenetic extension considered. Low current climate seasonalities were the main drivers of species richness and phylogenetic diversity variation across the Atlantic Forest. Whereas in higher phylogenetic extension, topographic heterogeneity increased the number of tree species independent of the sector, deep-past climate stability favored phylogenetic diversity by increasing relict lineages of distant clades in all forests, but with anomalies in the southern sector. This investigation yields substantial evidence that the response of the northern and southern sectors of the Atlantic Forest to identical environmental conditions diverges significantly, providing compelling support for the imprint of phylogenetic heritage in generating non-linear diversity patterns.

Trees structure the Earth's most biodiverse ecosystem, tropical forests. The vast number of tree species presents a formidable challenge to understanding these forests, including their response to environmental change, as very little is known about most tropical tree species. A focus on the common species may circumvent this challenge. Here we investigate abundance patterns of common tree species using inventory data on 1,003,805 trees with trunk diameters of at least 10 cm across 1,568 locations 1-6 in closed-canopy, structurally intact old-growth tropical forests in Africa, Amazonia and Southeast Asia. We estimate that 2.2%, 2.2% and 2.3% of species comprise 50% of the tropical trees in these regions, respectively. Extrapolating across all closed-canopy tropical forests, we estimate that just 1,053 species comprise half of Earth's 800 billion tropical trees with trunk diameters of at least 10 cm. Despite differing biogeographic, climatic and anthropogenic histories 7 , we find notably consistent patterns of common species and species abundance distributions across the continents. This suggests that fundamental mechanisms of tree community assembly may apply to all tropical forests. Resampling analyses show that the most common species are likely to belong to a manageable list of known species, enabling targeted efforts to understand their ecology. Although they do not detract from the importance of rare species, our results open new opportunities to understand the world's most diverse forests, including modelling their response to environmental change, by focusing on the common species that constitute the majority of their trees., (© 2024. The Author(s).)

Tropical tree diversity increases with rainfall 1,2 . Direct physiological effects of moisture availability and indirect effects mediated by biotic interactions are hypothesized to contribute to this pantropical increase in diversity with rainfall 2-6 . Previous studies have demonstrated direct physiological effects of variation in moisture availability on tree survival and diversity 5,7-10 , but the indirect effects of variation in moisture availability on diversity mediated by biotic interactions have not been shown 11 . Here we evaluate the relationships between interannual variation in moisture availability, the strength of density-dependent interactions, and seedling diversity in central Panama. Diversity increased with soil moisture over the first year of life across 20 annual cohorts. These first-year changes in diversity persisted for at least 15 years. Differential survival of moisture-sensitive species did not contribute to the observed changes in diversity. Rather, negative density-dependent interactions among conspecifics were stronger and increased diversity in wetter years. This suggests that moisture availability enhances diversity indirectly through moisture-sensitive, density-dependent conspecific interactions. Pathogens and phytophagous insects mediate interactions among seedlings in tropical forests 12-18 , and many of these plant enemies are themselves moisture-sensitive 19-27 . Changes in moisture availability caused by climate change and habitat degradation may alter these interactions and tropical tree diversity., (© 2023. The Author(s), under exclusive licence to Springer Nature Limited.)

Recent observations suggest that the large carbon sink in mature and recovering forests may be strongly limited by nitrogen 1-3 . Nitrogen-fixing trees (fixers) in symbiosis with bacteria provide the main natural source of new nitrogen to tropical forests 3,4 . However, abundances of fixers are tightly constrained 5-7 , highlighting the fundamental unanswered question of what limits new nitrogen entering tropical ecosystems. Here we examine whether herbivory by animals is responsible for limiting symbiotic nitrogen fixation in tropical forests. We evaluate whether nitrogen-fixing trees experience more herbivory than other trees, whether herbivory carries a substantial carbon cost, and whether high herbivory is a result of herbivores targeting the nitrogen-rich leaves of fixers 8,9 . We analysed 1,626 leaves from 350 seedlings of 43 tropical tree species in Panama and found that: (1) although herbivory reduces the growth and survival of all seedlings, nitrogen-fixing trees undergo 26% more herbivory than non-fixers; (2) fixers have 34% higher carbon opportunity costs owing to herbivory than non-fixers, exceeding the metabolic cost of fixing nitrogen; and (3) the high herbivory of fixers is not driven by high leaf nitrogen. Our findings reveal that herbivory may be sufficient to limit tropical symbiotic nitrogen fixation and could constrain its role in alleviating nitrogen limitation on the tropical carbon sink., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

Evidence exists that tree mortality is accelerating in some regions of the tropics 1,2 , with profound consequences for the future of the tropical carbon sink and the global anthropogenic carbon budget left to limit peak global warming below 2 °C. However, the mechanisms that may be driving such mortality changes and whether particular species are especially vulnerable remain unclear 3-8 . Here we analyse a 49-year record of tree dynamics from 24 old-growth forest plots encompassing a broad climatic gradient across the Australian moist tropics and find that annual tree mortality risk has, on average, doubled across all plots and species over the last 35 years, indicating a potential halving in life expectancy and carbon residence time. Associated losses in biomass were not offset by gains from growth and recruitment. Plots in less moist local climates presented higher average mortality risk, but local mean climate did not predict the pace of temporal increase in mortality risk. Species varied in the trajectories of their mortality risk, with the highest average risk found nearer to the upper end of the atmospheric vapour pressure deficit niches of species. A long-term increase in vapour pressure deficit was evident across the region, suggesting that thresholds involving atmospheric water stress, driven by global warming, may be a primary cause of increasing tree mortality in moist tropical forests., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

Natural disturbances are essential for tropical forests biodiversity. In the Afrotropics, megaherbivores have played a key role before their recent decline. Contrastingly to savanna elephants, forest elephants' impact on ecosystems remains poorly studied. Few decades ago, forests on Mount Cameroon were divided by lava flows, not being crossed by a local population of forest elephants until now. We assessed communities of trees, butterflies and two guilds of moths in the disturbed and undisturbed forests split by the longest lava flow. We surveyed 32 plots, recording 2025 trees of 97 species, and 7853 insects of 437 species. The disturbed forests differed in reduced tree density, height, and high canopy cover, and in increased DBH. Forest elephants' selective browsing and foraging also decreased tree species richness and altered their composition. The elephant disturbance increased butterfly species richness and had various effects on species richness and composition of the insect groups. These changes were likely caused by disturbance-driven alterations of habitats and species composition of trees. Moreover, the abandonment of forests by elephants led to local declines of range-restricted butterflies. The recent declines of forest elephants across the Afrotropics probably caused similar changes in forest biodiversity and should be reflected by conservation actions.

The flora of the Neotropics is unmatched in its diversity, however the mechanisms by which diversity has accumulated are debated and largely unclear. The Brownea clade (Leguminosae) is a characteristic component of the Neotropical flora, and the species within it are diverse in their floral morphology, attracting a wide variety of pollinators. This investigation aimed to estimate species divergence times and infer relationships within the group, in order to test whether the Brownea clade followed the 'cradle' or 'museum' model of diversification, i.e. whether species evolved rapidly over a short time period, or gradually over many millions of years. We also aimed to trace the spatio-temporal evolution of the clade by estimating ancestral biogeographical patterns in the group. We used BEAST to build a dated phylogeny of 73 Brownea clade species using three molecular markers (ITS, trnK and psbA-trnH), resulting in well-resolved phylogenetic relationships within the clade, as well as robust divergence time estimates from which we inferred diversification rates and ancestral biogeography. Our analyses revealed an Eocene origin for the group, after which the majority of diversification happened in Amazonia during the Miocene, most likely concurrent with climatic and geological changes caused by the rise of the Andes. We found no shifts in diversification rate over time, suggesting a gradual accumulation of lineages with low extinction rates. These results may help to understand why Amazonia is host to the highest diversity of tree species on Earth., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Although fungi play essential roles in nutrient cycles and plant growth in forest ecosystems, limited information is currently available on the community compositions of soil fungi in tropical forests. Few studies have examined fungal community structures in seasonal tropical forests, in which forest fires potentially have a large impact on above- and belowground community processes. Based on high-throughput sequencing technologies, we herein examined the diversity and community structures of soil fungi in dry seasonal tropical forests in Sakaerat, northeast Thailand. We found that fungal community compositions diverged among dry evergreen, dry deciduous, and fire-protected dry deciduous forests within the region. Although tree species diversity did not positively correlate with soil fungal diversity, the coverage of an understory bamboo species (Vietnamosasa pusilla) showed a strong relationship with fungal community structures. Our community ecological analysis also yielded a list of fungi showing habitat preferences for either of the neighboring evergreen and deciduous forests in Sakaerat. The present results provide a basis for managing soil fungal communities and aboveground plant communities in seasonal tropical forests in Southeast Asia.

Phosphorus availability is widely assumed to limit primary productivity in tropical forests, but support for this paradigm is equivocal. Although biogeochemical theory predicts that phosphorus limitation should be prevalent on old, strongly weathered soils, experimental manipulations have failed to detect a consistent response to phosphorus addition in species-rich lowland tropical forests. Here we show, by quantifying the growth of 541 tropical tree species across a steep natural phosphorus gradient in Panama, that phosphorus limitation is widespread at the level of individual species and strengthens markedly below a threshold of two parts per million exchangeable soil phosphate. However, this pervasive species-specific phosphorus limitation does not translate into a community-wide response, because some species grow rapidly on infertile soils despite extremely low phosphorus availability. These results redefine our understanding of nutrient limitation in diverse plant communities and have important implications for attempts to predict the response of tropical forests to environmental change.

Understanding the structure and dynamics of highly diverse tropical forests is challenging. Here we investigate the factors that drive the spatio-temporal variation of local tree numbers and species richness in a tropical forest (including 1250 plots of 20 × 20 m 2 ). To this end, we use a series of dynamic models that are built around the local spatial variation of mortality and recruitment rates, and ask which combination of processes can explain the observed spatial and temporal variation in tree and species numbers. We find that processes not included in classical neutral theory are needed to explain these fundamental patterns of the observed local forest dynamics. We identified a large spatio-temporal variability in the local number of recruits as the main missing mechanism, whereas variability of mortality rates contributed to a lesser extent. We also found that local tree numbers stabilize at typical values which can be explained by a simple analytical model. Our study emphasized the importance of spatio-temporal variability in recruitment beyond demographic stochasticity for explaining the local heterogeneity of tropical forests., (© 2017 The Author(s).)

Fossil and phylogenetic evidence indicates legume-rich modern tropical forests replaced Late Cretaceous palm-dominated tropical forests across four continents during the early Cenozoic (58-42 Ma). Tropical legume trees can transform ecosystems via their ability to fix dinitrogen (N 2 ) and higher leaf N compared with non-legumes (35-65%), but it is unclear how their evolutionary rise contributed to silicate weathering, the long-term sink for atmospheric carbon dioxide (CO 2 ). Here we hypothesize that the increasing abundance of N 2 -fixing legumes in tropical forests amplified silicate weathering rates by increased input of fixed nitrogen (N) to terrestrial ecosystems via interrelated mechanisms including increasing microbial respiration and soil acidification, and stimulating forest net primary productivity. We suggest the high CO 2 early Cenozoic atmosphere further amplified legume weathering. Evolution of legumes with high weathering rates was probably driven by their high demand for phosphorus and micronutrients required for N 2 -fixation and nodule formation., (© 2017 The Author(s).)

Currently, there is a lack of studies on the correct utilization of continuous distributions for dry tropical forests. Therefore, this work aims to investigate the diameter structure of a brazilian tropical dry forest and to select suitable continuous distributions by means of statistic tools for the stand and the main species. Two subsets were randomly selected from 40 plots. Diameter at base height was obtained. The following functions were tested: log-normal; gamma; Weibull 2P and Burr. The best fits were selected by Akaike's information validation criterion. Overall, the diameter distribution of the dry tropical forest was better described by negative exponential curves and positive skewness. The forest studied showed diameter distributions with decreasing probability for larger trees. This behavior was observed for both the main species and the stand. The generalization of the function fitted for the main species show that the development of individual models is needed. The Burr function showed good flexibility to describe the diameter structure of the stand and the behavior of Mimosa ophthalmocentra and Bauhinia cheilantha species. For Poincianella bracteosa, Aspidosperma pyrifolium and Myracrodum urundeuva better fitting was obtained with the log-normal function.

Lineages tend to retain ecological characteristics of their ancestors through time. However, for some traits, selection during evolutionary history may have also played a role in determining trait values. To address the relative importance of these processes requires large-scale quantification of traits and evolutionary relationships among species. The Amazonian tree flora comprises a high diversity of angiosperm lineages and species with widely differing life-history characteristics, providing an excellent system to investigate the combined influences of evolutionary heritage and selection in determining trait variation. We used trait data related to the major axes of life-history variation among tropical trees (e.g. growth and mortality rates) from 577 inventory plots in closed-canopy forest, mapped onto a phylogenetic hypothesis spanning more than 300 genera including all major angiosperm clades to test for evolutionary constraints on traits. We found significant phylogenetic signal (PS) for all traits, consistent with evolutionarily related genera having more similar characteristics than expected by chance. Although there is also evidence for repeated evolution of pioneer and shade tolerant life-history strategies within independent lineages, the existence of significant PS allows clearer predictions of the links between evolutionary diversity, ecosystem function and the response of tropical forests to global change., (© 2016 The Authors.)

Based on three 1-hm 2 plots of Jianfengling tropical montane rainforest on Hainan Island, 11 commom used functional traits of canopy trees were measured. After combining with topographical factors and trees census data of these three plots, we compared the impacts of weighted species abundance on two functional dispersion indices, mean pairwise distance (MPD) and mean nearest taxon distance (MNTD), by using single- and multi-dimensional traits, respectively. The relationship between functional richness of the forest canopies and species abundance was analyzed. We used a null model approach to explore the variations in standardized size effects of MPD and MNTD, which were weighted by species abundance and eliminated the influences of species richness diffe-rences among communities, and assessed functional diversity patterns of the forest canopies and their responses to local habitat heterogeneity at community's level. The results showed that variation in MPD was greatly dependent on the dimensionalities of functional traits as well as species abundance. The correlations between weighted and non-weighted MPD based on different dimensional traits were relatively weak (R=0.359-0.628). On the contrary, functional traits and species abundance had relatively weak effects on MNTD, which brought stronger correlations between weighted and non-weighted MNTD based on different dimensional traits (R=0.746-0.820). Functional dispersion of the forest canopies were generally overestimated when using non-weighted MPD and MNTD. Functional richness of the forest canopies showed an exponential relationship with species abundance (F=128.20; R 2 =0.632; AIC=97.72; P＜0.001), which might exist a species abundance threshold value. Patterns of functional diversity of the forest canopies based on different dimensional functional traits and their habitat responses showed variations in some degree. Forest canopies in the valley usually had relatively stronger biological competition, and functional diversity was higher than expected functional diversity randomized by null model, which indicated dispersed distribution of functional traits among canopy tree species in this habitat. However, the functional diversity of the forest canopies tended to be close or lower than randomization in the other habitat types, which demonstrated random or clustered distribution of the functional traits among canopy tree species.

Studies of forest dynamics plots (FDPs) have revealed a variety of negative density-dependent (NDD) demographic interactions, especially among conspecific trees. These interactions can affect growth rate, recruitment and mortality, and they play a central role in the maintenance of species diversity in these complex ecosystems. Here we use an equal area annulus (EAA) point-pattern method to comprehensively analyze data from two tropical FDPs, Barro Colorado Island in Panama and Sinharaja in Sri Lanka. We show that these NDD interactions also influence the continued evolutionary diversification of even distantly related tree species in these FDPs. We examine the details of a wide range of these interactions between individual trees and the trees that surround them. All these interactions, and their cumulative effects, are strongest among conspecific focal and surrounding tree species in both FDPs. They diminish in magnitude with increasing phylogenetic distance between heterospecific focal and surrounding trees, but do not disappear or change the pattern of their dependence on size, density, frequency or physical distance even among the most distantly related trees. The phylogenetic persistence of all these effects provides evidence that interactions between tree species that share an ecosystem may continue to promote adaptive divergence even after the species' gene pools have become separated. Adaptive divergence among taxa would operate in stark contrast to an alternative possibility that has previously been suggested, that distantly related species with dispersal-limited distributions and confronted with unpredictable neighbors will tend to converge on common strategies of resource use. In addition, we have also uncovered a positive density-dependent effect: growth rates of large trees are boosted in the presence of a smaller basal area of surrounding trees. We also show that many of the NDD interactions switch sign rapidly as focal trees grow in size, and that their cumulative effect can strongly influence the distributions and species composition of the trees that surround the focal trees during the focal trees' lifetimes., Competing Interests: The authors have declared that no competing interests exist.

The functional biogeography of tropical forests is expressed in foliar chemicals that are key physiologically based predictors of plant adaptation to changing environmental conditions including climate. However, understanding the degree to which environmental filters sort the canopy chemical characteristics of forest canopies remains a challenge. Here, we report on the elevation and soil-type dependence of forest canopy chemistry among 75 compositionally and environmentally distinct forests in nine regions, with a total of 7819 individual trees representing 3246 species collected, identified and assayed for foliar traits. We assessed whether there are consistent relationships between canopy chemical traits and both elevation and soil type, and evaluated the general role of phylogeny in mediating patterns of canopy traits within and across communities. Chemical trait variation and partitioning suggested a general model based on four interconnected findings. First, geographic variation at the soil-Order level, expressing broad changes in fertility, underpins major shifts in foliar phosphorus (P) and calcium (Ca). Second, elevation-dependent shifts in average community leaf dry mass per area (LMA), chlorophyll, and carbon allocation (including nonstructural carbohydrates) are most strongly correlated with changes in foliar Ca. Third, chemical diversity within communities is driven by differences between species rather than by plasticity within species. Finally, elevation- and soil-dependent changes in N, LMA and leaf carbon allocation are mediated by canopy compositional turnover, whereas foliar P and Ca are driven more by changes in site conditions than by phylogeny. Our findings have broad implications for understanding the global ecology of humid tropical forests, and their functional responses to changing climate., (© 2015 John Wiley & Sons Ltd.)

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 = 0.31-0.79). For all 12 focal species, most of the leaf litter fell less than 5 m away from the source trees, generating fine- scale 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.

There is an increasing interest in identifying theories, empirical data sets, and remote-sensing metrics that can quantify tropical forest alpha diversity at a landscape scale. Quantifying patterns of tree species richness in the field is time consuming, especially in regions with over 100 tree species/ha. We examine species richness in a 50-ha plot in Barro Colorado Island in Panama and test if biophysical measurements of canopy reflectance from high-resolution satellite imagery and detailed vertical forest structure and topography from light detection and ranging (lidar) are associated with species richness across four tree size classes (>1, 1-10, >10, and >20 cm dbh) and three spatial scales (1, 0.25, and 0.04 ha). We use the 2010 tree inventory, including 204,757 individuals belonging to 301 species of freestanding woody plants or 166 ± 1.5 species/ha (mean ± SE), to compare with remote-sensing data. All remote-sensing metrics became less correlated with species richness as spatial resolution decreased from 1.0 ha to 0.04 ha and tree size increased from 1 cm to 20 cm dbh. When all stems with dbh > 1 cm in 1-ha plots were compared to remote-sensing metrics, standard deviation in canopy reflectance explained 13% of the variance in species richness. The standard deviations of canopy height and the topographic wetness index (TWI) derived from lidar were the best metrics to explain the spatial variance in species richness (15% and 24%, respectively). Using multiple regression models, we made predictions of species richness across Barro Colorado Island (BCI) at the 1-ha spatial scale for different tree size classes. We predicted variation in tree species richness among all plants (adjusted r² = 0.35) and trees with dbh > 10 cm (adjusted r² = 0.25). However, the best model results were for understory trees and shrubs (dbh 1-10 cm) (adjusted r² = 0.52) that comprise the majority of species richness in tropical forests. Our results indicate that high-resolution remote sensing can predict a large percentage of variance in species richness and potentially provide a framework to map and predict alpha diversity among trees in diverse tropical forests.

The high species richness of tropical forests has long been recognized, yet there remains substantial uncertainty regarding the actual number of tropical tree species. Using a pantropical tree inventory database from closed canopy forests, consisting of 657,630 trees belonging to 11,371 species, we use a fitted value of Fisher's alpha and an approximate pantropical stem total to estimate the minimum number of tropical forest tree species to fall between ∼ 40,000 and ∼ 53,000, i.e., at the high end of previous estimates. Contrary to common assumption, the Indo-Pacific region was found to be as species-rich as the Neotropics, with both regions having a minimum of ∼ 19,000-25,000 tree species. Continental Africa is relatively depauperate with a minimum of ∼ 4,500-6,000 tree species. Very few species are shared among the African, American, and the Indo-Pacific regions. We provide a methodological framework for estimating species richness in trees that may help refine species richness estimates of tree-dependent taxa.

Understanding nutrient limitation of net primary productivity (NPP) is critical to predict how plant communities will respond to environmental change. Foliar nutrients, especially nitrogen and phosphorus concentrations ([N] and [P]) and their ratio, have been used widely as indicators of plant nutritional status and have been linked directly to nutrient limitation of NPP. In tropical systems, however, a high number of confounding factors can limit the ability to predict nutrient limitation--as defined mechanistically by NPP responses to fertilization--based on the stoichiometric signal of the plant community. We used a long-term full factorial N and P fertilization experiment in a lowland tropical wet forest in Costa Rica to explore how tissue (foliar, litter and root) [N] and [P] changed with fertilization, how different tree size classes and taxa influenced the community response, and how tissue nutrients related to NPP. Consistent with NPP responses to fertilization, there were no changes in community-wide foliar [N] and [P], two years after fertilization. Nevertheless, litterfall [N] increased with N additions and root [P] increased with P additions. The most common tree species (Pentaclethra macroloba) had 9% higher mean foliar [N] with NP additions and the most common palm species (Socratea exohrriza) had 15% and 19% higher mean foliar [P] with P and NP additions, respectively. Moreover, N:P ratios were not indicative of NPP responses to fertilization, either at the community or at the taxa level. Our study suggests that in these diverse tropical forests, tissue [N] and [P] are driven by the interaction of multiple factors and are not always indicative of the nutritional status of the plant community.

Background: India is rich with biodiversity, which includes a large number of endemic, rare and threatened plant species. Previous studies have used DNA barcoding to inventory species for applications in biodiversity monitoring, conservation impact assessment, monitoring of illegal trading, authentication of traded medicinal plants etc. This is the first tropical dry evergreen forest (TDEF) barcode study in the World and the first attempt to assemble a reference barcode library for the trees of India as part of a larger project initiated by this research group., Methodology/principal Findings: We sampled 429 trees representing 143 tropical dry evergreen forest (TDEF) species, which included 16 threatened species. DNA barcoding was completed using rbcL and matK markers. The tiered approach (1st tier rbcL; 2nd tier matK) correctly identified 136 out of 143 species (95%). This high level of species resolution was largely due to the fact that the tree species were taxonomically diverse in the TDEF. Ability to resolve taxonomically diverse tree species of TDEF was comparable among the best match method, the phylogenetic method, and the characteristic attribute organization system method., Conclusions: We demonstrated the utility of the TDEF reference barcode library to authenticate wood samples from timber operations in the TDEF. This pilot research study will enable more comprehensive surveys of the illegal timber trade of threatened species in the TDEF. This TDEF reference barcode library also contains trees that have medicinal properties, which could be used to monitor unsustainable and indiscriminate collection of plants from the wild for their medicinal value.

The integration of ecology and evolutionary biology requires an understanding of the evolutionary lability in species' ecological niches. For tropical trees, specialization for particular soil resource and topographic conditions is an important part of the habitat niche, influencing the distributions of individual species and overall tree community structure at the local scale. However, little is known about how these habitat niches are related to the evolutionary history of species. We assessed the relationship between taxonomic rank and tree species' soil resource and topographic niches in eight large (24-50 ha) tropical forest dynamics plots. Niche overlap values, indicating the similarity of two species' distributions along soil or topographic axes, were calculated for all pairwise combinations of co-occurring tree species at each study site. Congeneric species pairs often showed greater niche overlap (i.e., more similar niches) than non-congeneric pairs along both soil and topographic axes, though significant effects were found for only five sites based on Mantel tests. No evidence for taxonomic effects was found at the family level. Our results indicate that local habitat niches of trees exhibit varying degrees of phylogenetic signal at different sites, which may have important ramifications for the phylogenetic structure of these communities.

Two main theories have attempted to explain variation in plant species composition (β-diversity). Niche theory proposes that most of the variation is related to environment (environmental filtering), whereas neutral theory posits that dispersal limitation is the main driver of β-diversity. In this study, we first explored how α- and β-diversity of plant functional groups defined by growth form (trees, shrubs and lianas, which represent different strategies of resource partitioning), and dispersal syndrome (autochory, anemochory and zoochory, which represent differences in dispersal limitation) vary with successional age and topographic position in a tropical dry forest. Second, we examined the effects of environmental, spatial, and spatially-structured environmental factors on β-diversity of functional groups; we used the spatial structure of sampling sites as a proxy for dispersal limitation, and elevation, soil properties and forest stand age as indicators of environmental filtering. We recorded 200 species and 22,245 individuals in 276 plots; 120 species were trees, 41 shrubs and 39 lianas. We found that β-diversity was highest for shrubs, intermediate for lianas and lowest for trees, and was slightly higher for zoochorous than for autochorous and anemochorous species. All three dispersal syndromes, trees and shrubs varied in composition among vegetation classes (successional age and topographic position), whilst lianas did not. β-diversity was influenced mostly by proxies of environmental filtering, except for shrubs, for which the influence of dispersal limitation was more important. Stand age and topography significantly influenced α-diversity across functional groups, but showed a low influence on β-diversity -possibly due to the counterbalancing effect of resprouting on plant distribution and composition. Our results show that considering different plant functional groups reveals important differences in both α- and β-diversity patterns and correlates that are not apparent when focusing on overall woody plant diversity, and that have important implications for ecological theory and biodiversity conservation.

Background: DNA barcoding as a tool for species identification has been successful in animals and other organisms, including certain groups of plants. The exploration of this new tool for species identification, particularly in tree species, is very scanty from biodiversity-rich countries like India. rbcL and matK are standard barcode loci while ITS, and trnH-psbA are considered as supplementary loci for plants., Methodology and Principal Findings: Plant barcode loci, namely, rbcL, matK, ITS, trnH-psbA, and the recently proposed ITS2, were tested for their efficacy as barcode loci using 300 accessions of tropical tree species. We tested these loci for PCR, sequencing success, and species discrimination ability using three methods. rbcL was the best locus as far as PCR and sequencing success rate were concerned, but not for the species discrimination ability of tropical tree species. ITS and trnH-psbA were the second best loci in PCR and sequencing success, respectively. The species discrimination ability of ITS ranged from 24.4 percent to 74.3 percent and that of trnH-psbA was 25.6 percent to 67.7 percent, depending upon the data set and the method used. matK provided the least PCR success, followed by ITS2 (59. 0%). Species resolution by ITS2 and rbcL ranged from 9.0 percent to 48.7 percent and 13.2 percent to 43.6 percent, respectively. Further, we observed that the NCBI nucleotide database is poorly represented by the sequences of barcode loci studied here for tree species., Conclusion: Although a conservative approach of a success rate of 60-70 percent by both ITS and trnH-psbA may not be considered as highly successful but would certainly help in large-scale biodiversity inventorization, particularly for tropical tree species, considering the standard success rate of plant DNA barcode program reported so far. The recommended matK and rbcL primers combination may not work in tropical tree species as barcode markers.

Relationships between functional traits and average or potential demographic rates have provided insight into the functional constraints and trade-offs underlying life-history strategies of tropical tree species. We have extended this framework by decomposing growth rates of -130 000 trees of 171 Neotropical tree species into intrinsic growth and the response of growth to light and size. We related these growth characteristics to multiple functional traits (wood density, adult stature, seed mass, leaf traits) in a hierarchical Bayesian model that accounted for measurement error and intraspecific variability of functional traits. Wood density was the most important trait determining all three growth characteristics. Intrinsic growth rates were additionally strongly related to adult stature, while all traits contributed to light response. Our analysis yielded a predictive model that allows estimation of growth characteristics for rare species on the basis of a few easily measurable morphological traits.

1. Species diversity of arboreal arthropods tends to increase during rainforest succession so that primary forest communities comprise more species than those from secondary vegetation, but it is not well understood why. Primary forests differ from secondary forests in a wide array of factors whose relative impacts on arthropod diversity have not yet been quantified. 2. We assessed the effects of succession-related determinants on a keystone ecological group, arboreal ants, by conducting a complete census of 1332 ant nests from all trees with diameter at breast height ≥ 5 cm occurring within two (unreplicated) 0·32-ha plots, one in primary and one in secondary lowland forest in New Guinea. Specifically, we used a novel rarefaction-based approach to match number, size distribution and taxonomic structure of trees in primary forest communities to those in secondary forest and compared the resulting numbers of ant species. 3. In total, we recorded 80 nesting ant species from 389 trees in primary forest but only 42 species from 295 trees in secondary forest. The two habitats did not differ in the mean number of ant species per tree or in the relationship between ant diversity and tree size. However, the between-tree similarity of ant communities was higher in secondary forest than in primary forest, as was the between-tree nest site similarity, suggesting that secondary trees were more uniform in providing nesting microhabitats. 4. Using our rarefaction method, the difference in ant species richness between two forest types was partitioned according to the effects of higher tree density (22·6%), larger tree size (15·5%) and higher taxonomic diversity of trees (14·3%) in primary than in secondary forest. The remaining difference (47·6%) was because of higher beta diversity of ant communities between primary forest trees. In contrast, difference in nest density was explained solely by difference in tree density. 5. Our study shows that reduction in plant taxonomic diversity in secondary forests is not the main driver of the reduction in canopy ant species richness. We suggest that the majority of arboreal species losses in secondary tropical forests are attributable to simpler vegetation structure, combined with lower turnover of nesting microhabitats between trees., (© 2012 The Authors. Journal of Animal Ecology © 2012 British Ecological Society.)

Plant diversity is considered one factor structuring soil fungal communities because the diversity of compounds in leaf litter might determine the extent of resource heterogeneity for decomposer communities. Lowland tropical rain forests have the highest plant diversity per area of any biome. Since fungi are responsible for much of the decomposition occurring in forest soils, understanding the factors that structure fungi in tropical forests may provide valuable insight for predicting changes in global carbon and nitrogen fluxes. To test the role of plant diversity in shaping fungal community structure and function, soil (0-20 cm) and leaf litter (O horizons) were collected from six established 1-ha forest census plots across a natural plant diversity gradient on the Isthmus of Panama. We used 454 pyrosequencing and phospholipid fatty acid analysis to evaluate correlations between microbial community composition, precipitation, soil nutrients, and plant richness. In soil, the number of fungal taxa increased significantly with increasing mean annual precipitation, but not with plant richness. There were no correlations between fungal communities in leaf litter and plant diversity or precipitation, and fungal communities were found to be compositionally distinct between soil and leaf litter. To directly test for effects of plant species richness on fungal diversity and function, we experimentally re-created litter diversity gradients in litter bags with 1, 25, and 50 species of litter. After 6 months, we found a significant effect of litter diversity on decomposition rate between one and 25 species of leaf litter. However, fungal richness did not track plant species richness. Although studies in a broader range of sites is required, these results suggest that precipitation may be a more important factor than plant diversity or soil nutrient status in structuring tropical forest soil fungal communities.

The degree to which turnover in biological communities is structured by deterministic or stochastic factors and the identities of influential deterministic factors are fundamental, yet unresolved, questions in ecology. Answers to these questions are particularly important for projecting the fate of forests with diverse disturbance histories worldwide. To uncover the processes governing turnover we use species-level molecular phylogenies and functional trait data sets for two long-term tropical forest plots with contrasting disturbance histories: one forest is older-growth, and one was recently disturbed. Having both phylogenetic and functional information further allows us to parse out the deterministic influences of different ecological filters. With the use of null models we find that compositional turnover was random with respect to phylogeny on average, but highly nonrandom with respect to measured functional traits. Furthermore, as predicted by a deterministic assembly process, the older-growth and disturbed forests were characterized by less than and greater than expected functional turnover, respectively. These results suggest that the abiotic environment, which changes due to succession in the disturbed forest, strongly governs the temporal dynamics of disturbed and undisturbed tropical forests. Predicting future changes in the composition of disturbed and undisturbed forests may therefore be tractable when using a functional-trait-based approach.

The SITE® model was originally developed to study the response of tropical ecosystems to varying environmental conditions. The present study evaluated the applicability of the SITE model to simulation of energy fluxes in a tropical semi-deciduous forest of the southern Amazon Basin. The model was simulated with data representing the wet and dry season, and was calibrated according to each season. The output data of the calibrated model [net radiation (Rn), latent heat flux (LE) and sensible heat flux (H)] were compared with data observed in the field for validation. Considering changes in parameter calibration for a time step simulation of 30 min, the magnitude of variation in temporal flux was satisfactory when compared to observation field data. There was a tendency to underestimate and overestimate LE and H, respectively. Of all the calibration parameters, the soil moisture parameter presented the highest variation over the seasons, thus influencing SITE model performance.

Leaf area index (LAI) is an important ecophysiological variable because leaves are the organs responsible for gas exchange between plants and the atmosphere. This variable can be calculated from primary values of leaf area assessed by destructive or non-destructive methods, which is relatively easy when crop species are investigated, but is not the case when the focus is on natural wood plants communities. In this paper, we analyze the seasonality of LAI estimated by three different methods in the Amazonia-savannah transitional forest, located 50 km north-east of Sinop city, Mato Grosso, Brazil. In the first method, we combine Monsi and Saekis' original method [Monsi M, Saeki T (1953) Jpn J Bot 14:22-52], which measures LAI using the Beer-Lambert extinction law, and the proposition of Goudriaan [Goudriaan J (1988) Agric For Meteorol 43:155-169] to estimate the extinction coefficient from solar height. The second method differed from the first only in the way in which the daily fraction of intercepted photosynthetic active radiation (FPAR) was calculated, as proposed by Charles-Edwards and Lawn (Charles-Edwards DA, Lawn RJ (1984) Plant Cell Environ 7:247-251]. In the third method, we used a remote sensing technique [MOD15&#95;BU-collection 4, produced and distributed by EROS Data Center Distributed Active Archive Center (EDC DAAC)]. We found that the first and the second methods revealed the expected LAI dynamics, which increased during the dry-wet transition and wet season, and decreased during the wet-dry transition and dry season. From 20 randomly distributed sets in a 1.0 ha area, only 3 showed significant differences in LAI estimated from the first two methods; conversely, LAI was overestimated by the third method.

Based on the investigation data of monsoon evergreen broad-leaved forest at its different succession stages (primary, CP; 15 years of succession, CF; and 30 years of succession, CT) in Pu' er of Yunnan Province, this paper studied the species-area relationship of this forest at each succession stage. It was found that in the communities at each succession stage, the number of total species, trees, shrubs, and lianas had a significant correlation with sampling area, with the area explained over 94% of the total variation. The Z value of the total species (0.334) and trees (0.394) was the lowest at CT, whereas that of shrubs (0.437) and lianas (0.326) was the lowest at CF. No significant differences were observed in the intercepts of the species-area curve of total species, trees, shrubs, and lianas among different succession stages, but the coefficient of determination (R2) of the species-area curve of total species and lianas was the highest at CP. The richness of trees and shrubs at CF explained 99.9% of the variance of Z value, but the richness of total species, trees, shrubs, and lianas at CP and CT had no significant correlations with the Z value.

Background: Plants interact with each other, nutrients, and microbial communities in soils through extensive root networks. Understanding these below ground interactions has been difficult in natural systems, particularly those with high plant species diversity where morphological identification of fine roots is difficult. We combine DNA-based root identification with a DNA barcode database and above ground stem locations in a floristically diverse lowland tropical wet forest on Barro Colorado Island, Panama, where all trees and lianas >1 cm diameter have been mapped to investigate richness patterns below ground and model rooting distributions., Methodology/principal Findings: DNA barcode loci, particularly the cpDNA locus trnH-psba, can be used to identify fine and small coarse roots to species. We recovered 33 species of roots from 117 fragments sequenced from 12 soil cores. Despite limited sampling, we recovered a high proportion of the known species in the focal hectare, representing approximately 14% of the measured woody plant richness. This high value is emphasized by the fact that we would need to sample on average 13 m(2) at the seedling layer and 45 m(2) for woody plants >1 cm diameter to obtain the same number of species above ground. Results from inverse models parameterized with the locations and sizes of adults and the species identifications of roots and sampling locations indicates a high potential for distal underground interactions among plants., Conclusions: DNA barcoding techniques coupled with modeling approaches should be broadly applicable to studying root distributions in any mapped vegetation plot. We discuss the implications of our results and outline how second-generation sequencing technology and environmental sampling can be combined to increase our understanding of how root distributions influence the potential for plant interactions in natural ecosystems.

Elucidating the ecological mechanisms underlying community assembly in subtropical forests remains a central challenge for ecologists. The assembly of species into communities can be due to interspecific differences in habitat associations, and there is increasing evidence that these associations may have an underlying phylogenetic structure in contemporary terrestrial communities. In other words, by examining the degree to which closely related species prefer similar habitats and the degree to which they co-occur, ecologists are able to infer the mechanisms underlying community assembly. Here we implement this approach in a diverse subtropical tree community in China using a long-term forest dynamics plot and a molecular phylogeny generated from three DNA barcode loci. We find that there is phylogenetic signal in plant-habitat associations (i.e. closely related species tend to prefer similar habitats) and that patterns of co-occurrence within habitats are typically non-random with respect to phylogeny. In particular, we found phylogenetic clustering in valley and low-slope habitats in this forest, indicating a filtering of lineages plays a dominant role in structuring communities in these habitats and we found evidence of phylogenetic overdispersion in high-slope, ridge-top and high-gully habitats, indicating that distantly related species tended to co-occur in these high elevation habitats and that lineage filtering is less important in structuring these communities. Thus we infer that non-neutral niche-based processes acting upon evolutionarily conserved habitat preferences explain the assembly of local scale communities in the forest studied.

Seasonally dry tropical forests (SDTF) are characterized by pronounced seasonality in rainfall, and as a result trees in these forests must endure seasonal variation in soil water availability. Furthermore, SDTF on the northern Yucatan Peninsula, Mexico, have a legacy of disturbances, thereby creating a patchy mosaic of different seral stages undergoing secondary succession. We examined the water status of six canopy tree species, representing contrasting leaf phenology (evergreen vs. drought-deciduous) at three seral stages along a fire chronosequence in order to better understand strategies that trees use to overcome seasonal water limitations. The early-seral forest was characterized by high soil water evaporation and low soil moisture, and consequently early-seral trees exhibited lower midday bulk leaf water potentials (Ψ(L)) relative to late-seral trees (-1.01 ± 0.14 and -0.54 ± 0.07 MPa, respectively). Although Ψ(L) did not differ between evergreen and drought-deciduous trees, results from stable isotope analyses indicated different strategies to overcome seasonal water limitations. Differences were especially pronounced in the early-seral stage where evergreen trees had significantly lower xylem water δ(18)O values relative to drought-deciduous trees (-2.6 ± 0.5 and 0.3 ± 0.6‰, respectively), indicating evergreen species used deeper sources of water. In contrast, drought-deciduous trees showed greater enrichment of foliar (18)O (∆(18)O(l)) and (13)C, suggesting lower stomatal conductance and greater water-use efficiency. Thus, the rapid development of deep roots appears to be an important strategy enabling evergreen species to overcome seasonal water limitation, whereas, in addition to losing a portion of their leaves, drought-deciduous trees minimize water loss from remaining leaves during the dry season.

Evergreen broad-leaved forest is now gradually degraded and fragmented, and there is an increase in the amount of habitat edges as a result of long-term human activity. However, the role of edges in the regeneration of primary forest species is poorly understood. After 20 years of the edge creation, we analyzed primary forest species distribution and abundance, and changes in floristic composition, vegetation structure across forest-field gradients in Ailao Mountain, SW China. Our results revealed that there was a higher abundance and richness of primary species, late secondary species and thorny lianas at the distances 0-50 m than at the distances more than 50 m from the edge into the forest exterior. At the distances >50 m, no individuals of dominant canopy trees Lithocarpus xylocarpus, Castanopsis wattii, and L. jingdongensis were found, whereas the abundance of early pioneer shrub species and herbaceous cover was significantly greater. The richness of primary species showed a decrease with increasing distances from the forest edge to the exterior, particularly of medium-seeded primary species showing a drastic decrease. Moreover, no large-seeded primary species occurred at the distances >60 m. This study indicates that the forest edge as a buffer zone may be in favor of primary species regeneration. A dense shrub and herb layer, and seed dispersal may be the major factors limiting the forest regeneration farther from the forest edge. Therefore, to facilitate forest recovery processes, management should give priority to the protection of buffer zones of this forest edge.

The accumulation of species-specific enemies around adults is hypothesized to maintain plant diversity by limiting the recruitment of conspecific seedlings relative to heterospecific seedlings. Although previous studies in forested ecosystems have documented patterns consistent with the process of negative feedback, these studies are unable to address which classes of enemies (for example, pathogens, invertebrates, mammals) exhibit species-specific effects strong enough to generate negative feedback, and whether negative feedback at the level of the individual tree is sufficient to influence community-wide forest composition. Here we use fully reciprocal shade-house and field experiments to test whether the performance of conspecific tree seedlings (relative to heterospecific seedlings) is reduced when grown in the presence of enemies associated with adult trees. Both experiments provide strong evidence for negative plant-soil feedback mediated by soil biota. In contrast, above-ground enemies (mammals, foliar herbivores and foliar pathogens) contributed little to negative feedback observed in the field. In both experiments, we found that tree species that showed stronger negative feedback were less common as adults in the forest community, indicating that susceptibility to soil biota may determine species relative abundance in these tropical forests. Finally, our simulation models confirm that the strength of local negative feedback that we measured is sufficient to produce the observed community-wide patterns in tree-species relative abundance. Our findings indicate that plant-soil feedback is an important mechanism that can maintain species diversity and explain patterns of tree-species relative abundance in tropical forests.

Lianas are an important component of tropical forests and often abundant in open habitats, such as tree-fall gaps, forest edges, and disturbed forests. The abundance of lianas in tropical forests has been increasing as a result of global environmental change and increasing forest fragmentation. In order to understand this phenomenon in terms of leaf functional traits and to evaluate their competitive potential, we conducted a cost-benefit analysis of leaves from 18 liana species and 19 tree species in a tropical seasonal rain forest. The results revealed that lianas were scattered in a group distinct from trees along the first axis of a principal component analysis using 15 leaf ecophysiological traits, being located at the quick-return end of the leaf economics spectrum, with higher specific leaf area and photosynthetic rates (A), higher photosynthetic nitrogen (N) and phosphorus (P) use efficiencies, a lower leaf construction cost per unit leaf area (CC) and cost-benefit ratio (CC/A), and a shorter leaf life span (LLS). Trees showed the opposite trends. The results indicate that lianas can grow faster and capture resources more efficiently than trees in disturbed, open habitats. The positive relationship between LLS and CC/A revealed a trade-off between leaf construction cost and benefit over time. The 37 species analyzed had a mean foliar N/P ratio of 20, indicating that the forest was characterized by a P deficit. With an increasing atmospheric CO(2) concentration, the higher nutrient use efficiency could benefit lianas more than trees in terms of productivity, possibly also contributing to the increasing abundance of lianas in nutrient-limited tropical forests.

Despite research demonstrating that water and nutrient availability exert strong effects on multiple ecosystem processes in tropical forests, little is known about the effect of these factors on the demography and population dynamics of tropical trees. Over the course of 5 years, we monitored two common Amazonian secondary forest species-Lacistema pubescens and Myrcia sylvatica-in dry-season irrigation, litter-removal and control plots. We then evaluated the effects of altered water and nutrient availability on population demography and dynamics using matrix models and life table response experiments. Our results show that despite prolonged experimental manipulation of water and nutrient availability, there were nearly no consistent and unidirectional treatment effects on the demography of either species. The patterns and significance of observed treatment effects were largely dependent on cross-year variability not related to rainfall patterns, and disappeared once we pooled data across years. Furthermore, most of these transient treatment effects had little effect on population growth rates. Our results suggest that despite major experimental manipulations of water and nutrient availability-factors considered critical to the ecology of tropical pioneer tree species-autogenic light limitation appears to be the primary regulator of tree demography at early/mid successional stages. Indeed, the effects of light availability may completely override those of other factors thought to influence the successional development of Amazonian secondary forests.

Resource availability and disturbance are important factors that shape the composition, structure, and functioning of ecosystems. We investigated the effects of soil fertility and disturbance on plant-soil interactions and nutrient cycling in a diverse tropical rainforest. Our goal was to determine how common soil specialisation is among species and how plant-soil interactions affect ecosystem functioning in the presence of disturbance. Most species (59%) showed significant fidelity to either fertile (basalt) or infertile (schist) soils. Obligate schist specialists (six species) contributed 39 and 37% to total stand-level basal area and aboveground net primary productivity, respectively. High nutrient use efficiency of schist specialists reduced the rates of within-stand nutrient cycling through the production of nutrient-poor plant tissues and litter. Although forests on schist soils had higher basal area and similar rates of productivity to forests on basalt, uptake of Mg, K, P, and N were markedly less on schist than on basalt, particularly after a cyclone disturbance. Stands on schist soils were also less affected by the cyclone and, as a result, contributed less (ca. 50%) Mg, K, P, and N inputs to the forest floor (via litterfall) than stands on basalt soils. System "openness" (i.e. the risk of nutrient loss) from cyclone-affected basalt forests was minimised by high rates of uptake following disturbance and large effective cation exchange capacities of soils. Soil-plant-disturbance interactions are likely to engender different fitness-enhancing strategies on fertile and infertile soils, possibly leading to the development and/or maintenance of diversity in rainforests.

The future of tropical forest biodiversity depends more than ever on the effective management of human-modified landscapes, presenting a daunting challenge to conservation practitioners and land use managers. We provide a critical synthesis of the scientific insights that guide our understanding of patterns and processes underpinning forest biodiversity in the human-modified tropics, and present a conceptual framework that integrates a broad range of social and ecological factors that define and contextualize the possible future of tropical forest species. A growing body of research demonstrates that spatial and temporal patterns of biodiversity are the dynamic product of interacting historical and contemporary human and ecological processes. These processes vary radically in their relative importance within and among regions, and have effects that may take years to become fully manifest. Interpreting biodiversity research findings is frequently made difficult by constrained study designs, low congruence in species responses to disturbance, shifting baselines and an over-dependence on comparative inferences from a small number of well studied localities. Spatial and temporal heterogeneity in the potential prospects for biodiversity conservation can be explained by regional differences in biotic vulnerability and anthropogenic legacies, an ever-tighter coupling of human-ecological systems and the influence of global environmental change. These differences provide both challenges and opportunities for biodiversity conservation. Building upon our synthesis we outline a simple adaptive-landscape planning framework that can help guide a new research agenda to enhance biodiversity conservation prospects in the human-modified tropics.

Today, the Seasonally Dry Tropical Forests (SDTF) of eastern South America occur as large, well-defined nuclei (e.g. Caatinga in the northeast) and as smaller enclaves within other vegetations (e.g. Cerrado and Chaco). In order to infer the way the present SDTF distribution was attained, the genetic structure of Astronium urundeuva, a tree confined to SDTF, was assessed using two chloroplast spacers and nine microsatellite loci. Five haplotypes were identified, whose distribution was spatially structured. The distribution of the two most common and divergent haplotypes suggested former vicariance and progressive divergence due to isolation. More recent range expansions of these two lineages subsequently occurred, leading to a secondary contact at the southern limit of the Caatinga SDTF nucleus. The multilocus-Bayesian approach using microsatellites consistently identified three groups of populations (Northeast, Central and Southwest). Isolation by distance was found in Northeast and Southwest groups whereas admixture was detected in the Central group, located at the transition between Caatinga and Cerrado domains. All together, the results support the existence of range expansions and secondary contact in the Central group. This study provides arguments that favour the existence of a previously more continuous formation of SDTF in eastern South America.

We investigated acclimation responses of seedlings and saplings of the pioneer species Cecropia schreberiana Miq. and three non-pioneer species, Dacryodes excelsa Vahl, Prestoea acuminata (Willdenow) H.E. Moore var. montana (Graham) Henderson and Galeano, and Sloanea berteriana Choisy ex DC, following a hurricane disturbance in a lower montane wet forest in Puerto Rico. Measurements were made, shortly after passage of the hurricane, on leaves expanded before the hurricane (pre-hurricane leaves) and, at a later time, on recently matured leaves that developed after the hurricane (post-hurricane leaves) from both seedlings and saplings at sites that were severely damaged by the hurricane (disturbed sites) and at sites with little disturbance (undisturbed sites). Pre-hurricane leaves of the non-pioneer species had relatively low light-saturated photosynthetic rates (A(max)) and stomatal conductance (g(s)); neither A(max) nor g(s) responded greatly to the increase in irradiance that resulted from the disturbance, and there were few significant differences between seedlings and saplings. Pre-hurricane leaves of plants at undisturbed sites had low dark respiration rates per unit area (R(d)) and light compensation points (LCP), whereas pre-hurricane leaves of plants at disturbed sites had significantly higher R(d) and LCP. Post-hurricane leaves of plants at disturbed sites had significantly higher A(max) and R(d) than plants at undisturbed sites. Compared with seedlings, saplings had higher A(max) and R(d) and showed greater acclimation to the increase in irradiance that followed the disturbance. Post-hurricane leaves of the non-pioneer species had significantly lower A(max) and were less responsive to changes in irradiance than the pioneer species C. schreberiana. Variation in A(max) across light environments and stages was strongly related to differences in leaf mass per unit area (LMA), especially in the non-pioneer species. As indicated by V(cmax) or J(max) per unit nitrogen, light acclimation of A(max) was determined by leaf morphology (LMA) for the non-pioneer species and by both leaf morphology and leaf biochemistry for C. schreberiana. Ontogenetic changes in A(max) were attributable to changes in leaf morphology. The ontogenetic component of variation in A(max) across light environments and stages differed among species, ranging from 36 to 59% for the non-pioneer species (D. excelsa, 59.3%; P. acuminata var. montana, 44.7%; and S. berteriana, 36.3%) compared with only 17% in the pioneer species C. schreberiana.

The contribution of mycorrhizal associations to maintaining tree diversity patterns in tropical rain forests is poorly known. Many tropical monodominant trees form ectomycorrhizal (EM) associations, and there is evidence that the EM mutualism contributes to the maintenance of monodominance. It is assumed that most other tropical tree species form arbuscular mycorrhizal (AM) associations, and while many mycorrhizal surveys have been done, the mycorrhizal status of numerous tropical tree taxa remains undocumented. In this study, we tested the assumption that most tropical trees form AM associations by sampling root vouchers from tree and liana species in monodominant Dicymbe corymbosa forest and an adjacent mixed rain forest in Guyana. Roots were assessed for the presence/ absence of AM and EM structures. Of the 142 species of trees and lianas surveyed, three tree species (the mono-dominant D. corymbosa, the grove-forming D. altsonii, and the non-dominant Aldina insignis) were EM, 137 were exclusively AM, and two were non-mycorrhizal. Both EM and AM structures wer e observed in D. corymbosa and D. altsonii. These results provide empirical data supporting the assumption that most tropical trees form AM associations for this region in the Guiana Shield and provide the first report of dual EM/AM colonization in Dicymbe species. Dual colonization of the Dicymbe species should be further explored to determine if this ability contributes to the establishment and maintenance of site dominance.