C. E. Timothy Paine

TREE RECRUITMENT IN AN EMPTY FOREST

To assess how the decimation of large vertebrates by hunting alters recruitment processes in a tropical forest, we compared the sapling cohorts of two structurally and compositionally similar forests in the Rio Manu floodplain in southeastern Peru. Large vertebrates were severely depleted at one site, Boca Manu (BM), whereas the other, Cocha Cashu Biological Station (CC), supported an intact fauna. At both sites we sampled small (> or =1 m tall, <1 cm dbh) and large (> or =1 cm and <10 cm dbh) saplings in the central portion of 4-ha plots within which all trees > or =10 cm dbh were mapped and identified. This design ensured that all conspecific adults within at least 50 m (BM) or 55 m (CC) of any sapling would have known locations. We used the Janzen-Connell model to make five predictions about the sapling cohorts at BM with respect to CC: (1) reduced overall sapling recruitment, (2) increased recruitment of species dispersed by abiotic means, (3) altered relative abundances of species, (4) prominence of large-seeded species among those showing depressed recruitment, and (5) little or no tendency for saplings to cluster closer to adults at BM. Our results affirmed each of these predictions. Interpreted at face value, the evidence suggests that few species are demographically stable at BM and that up to 28% are increasing and 72% decreasing. Loss of dispersal function allows species dispersed abiotically and by small birds and mammals to substitute for those dispersed by large birds and mammals. Although we regard these conclusions as preliminary, over the long run, the observed type of directional change in tree composition is likely to result in biodiversity loss and negative feedbacks on both the animal and plant communities. Our results suggest that the best, and perhaps only, way to prevent compositional change and probable loss of diversity in tropical tree communities is to prohibit hunting.

Rare species support vulnerable functions in high-diversity ecosystems

The most unusual, and thus irreplaceable, functions performed by species in three different species-rich ecosystems are fulfilled by only the rare species in these ecosystems.

Phylogenetic density dependence and environmental filtering predict seedling mortality in a tropical forest

Negative density dependence (NDD) and environmental filtering (EF) shape community assembly, but their relative importance is poorly understood. Recent studies have shown that seedlings mortality risk is positively related to the phylogenetic relatedness of neighbours. However, natural enemies, whose depredations often cause NDD, respond to functional traits of hosts rather than phylogenetic relatedness per se. To understand the roles of NDD and EF in community assembly, we assessed the effects on seedling mortality of functional similarity, phylogenetic relatedness and stem density of neighbouring seedlings and adults in a species-rich tropical forest. Mortality risks increased for common species when their functional traits departed substantially from the neighbourhood mean, and for all species when surrounded by close relatives. This indicates that NDD affects community assembly more broadly than does EF, and leads to the tentative conclusion that natural enemies respond to phylogenetically correlated traits. Our results affirm the prominence of NDD in structuring species-rich communities.

SEED PREDATION BY NEOTROPICAL RAIN FOREST MAMMALS INCREASES DIVERSITY IN SEEDLING RECRUITMENT

Seed dispersal and seedling recruitment (the transition of seeds to seedlings) set the spatiotemporal distribution of new individuals in plant communities. Many terrestrial rain forest mammals consume post-dispersal seeds and seedlings, often inflicting density-dependent mortality. In part because of density-dependent mortality, diversity often increases during seedling recruitment, making it a critical stage for species coexistence. We determined how mammalian predators, adult tree abundance, and seed mass interact to affect seedling recruitment in a western Amazonian rain forest. We used exclosures that were selectively permeable to three size classes of mammals: mice and spiny rats (weighing <1 kg), medium-sized rodents (1-12 kg), and large mammals (20-200 kg). Into each exclosure, we placed seeds of 13 tree species and one canopy liana, which varied by an order of magnitude in adult abundance and seed mass. We followed the fates of the seeds and resulting seedlings for at least 17 months. We assessed the effect of each mammalian size class on seed survival, seedling survival and growth, and the density and diversity of the seedlings that survived to the end of the experiment. Surprisingly, large mammals had no detectable effect at any stage of seedling recruitment. In contrast, small- and medium-sized mammals significantly reduced seed survival, seedling survival, and seedling density. Furthermore, predation by small mammals increased species richness on a per-stem basis. This increase in diversity resulted from their disproportionately intense predation on common species and large-seeded species. Small mammals thereby generated a rare-species advantage in seedling recruitment, the critical ingredient for frequency dependence. Predation by small (and to a lesser extent, medium-sized) mammals on seeds and seedlings significantly increases tree species diversity in tropical forests. This is the first long-term study to dissect the effects of various mammalian predators on the recruitment of a diverse set of tree species.

Environmental factors predict community functional composition in Amazonian forests

Summary 1. The consequences of biodiversity loss for ecosystem services largely depend on the functional identities of extirpated species. However, poor descriptions of spatial patterns of community functional composition across landscapes hamper accurate predictions, particularly in highly diverse tropical regions. Therefore, understanding how community functional composition varies across environmental gradients remains an important challenge. 2. We sampled 15 functional traits in 800 Neotropical tree species across 13 forest plots representative of the broad climatic and soil gradients encompassed by three widespread lowland forest habitats (terra firme forests on clay-rich soils, seasonally flooded forests and white-sand forests) at opposite ends of Amazonia (Peru and French Guiana). We combined univariate and multivariate approaches to test the magnitude and predictability of environmental filtering on community leaf and wood functional composition. 3. Directional shifts in community functional composition correlated with environmental changes across the 13 plots, with denser leaves, stems and roots in forests occurring in environments with limited water and soil-nutrient availability. Critically, these relationships allowed us to accurately predict the functional composition of 61 additional forest plots from environmental data alone. 4. Synthesis. Environmental filtering consistently shapes the functional composition of highly diverse tropical forests at large scales across the terra firme, seasonally flooded and white-sand forests of lowland Amazonia. Environmental factors drive and allow the prediction of variation in community functional composition among habitat types in Amazonian forests.

Continental-scale patterns of Cecropia reproductive phenology: evidence from herbarium specimens

Plant phenology is concerned with the timing of recurring biological events. Though phenology has traditionally been studied using intensive surveys of a local flora, results from such surveys are difficult to generalize to broader spatial scales. In this study, contrastingly, we assembled a continental-scale dataset of herbarium specimens for the emblematic genus of Neotropical pioneer trees, Cecropia, and applied Fourier spectral and cospectral analyses to investigate the reproductive phenology of 35 species. We detected significant annual, sub-annual and continuous patterns, and discuss the variation in patterns within and among climatic regions. Although previous studies have suggested that pioneer species generally produce flowers continually throughout the year, we found that at least one third of Cecropia species are characterized by clear annual flowering behaviour. We further investigated the relationships between phenology and climate seasonality, showing strong associations between phenology and seasonal variations in precipitation and temperature. We also verified our results against field survey data gathered from the literature. Our findings indicate that herbarium material is a reliable resource for use in the investigation of large-scale patterns in plant phenology, offering a promising complement to local intensive field studies.

spacodiR: structuring of phylogenetic diversity in ecological communities

MOTIVATION spacodiR is a cross-platform package, written for the R environment, for studying partitioning of diversity among natural communities in space and time. Complementing and extending existing software, spacodiR allows for hypothesis testing and parameter estimation in studying spatial structuring of species-, phylogenetic- and trait diversities. AVAILABILITY Integrated with other software in the R environment and with well documented and demonstrated functions, spacodiR is an open-source package and available at http://cran.r-project.org. CONTACT jonathan.eastman@gmail.com; ohardy@ulb.ac.be.

Within-individual variation of trunk and branch xylem density in tropical trees

PREMISE OF THE STUDY Wood density correlates with mechanical and physiological strategies of trees and is important for estimating global carbon stocks. Nonetheless, the relationship between branch and trunk xylem density has been poorly explored in neotropical trees. Here, we examine this relationship in trees from French Guiana and its variation among different families and sites, to improve the understanding of wood density in neotropical forests. METHODS Trunk and branch xylem densities were measured for 1909 trees in seven sites across French Guiana. A major-axis fit was performed to explore their general allometric relationship and its variation among different families and sites. KEY RESULTS Trunk xylem and branch xylem densities were significantly positively correlated, and their relationship explained 47% of the total variance. Trunk xylem was on average 9% denser than branch xylem. Family-level differences and interactions between family and site accounted for more than 40% of the total variance, whereas differences among sites explained little variation. CONCLUSIONS Variation in xylem density within individual trees can be substantial, and the relationship between branch xylem and trunk xylem densities varies considerably among families and sites. As such, whole-tree biomass estimates based on nondestructive branch sampling should correct for both taxonomic and environmental factors. Furthermore, detailed estimates of the vertical distribution of wood density within individual trees are needed to determine the extent to which relying solely upon measures of trunk wood density may cause carbon stocks in tropical forests to be overestimated.

Supplemental irrigation increases seedling performance and diversity in a tropical forest.

Diversity is positively correlated with water availability at global, continental and regional scales. With the objective of better understanding the mechanisms that drive these relationships, we investigated the degree to which variation in water availability affects the performance (recruitment, growth and survival) of juvenile trees. Precipitation was supplemented throughout two dry seasons in a seasonal moist forest in south-eastern Peru. Supplementing precipitation by 160 mm mo -1 , we increased soil moisture by 17%. To generate seedling communities of known species composition, we sowed 3840 seeds of 12 species. We monitored the fates of the 554 seedlings recruited from the sown seeds, as well as 1856 older non-sown seedlings (10 cm ≤ height 1 m tall). Watering significantly enhanced young seedling growth and survival, increasing stem density and diversity. Watering diminished the recruitment of species associated with upland forests, but increased the survival of both upland- and lowland-associated species. Though supplemental watering increased the growth of older seedlings, their density and diversity were unaffected. Sapling performance was insensitive to watering. We infer that variation in dry-season water availability may affect seedling community structure by differentially affecting recruitment and increasing overall survival. These results suggest that differential seedling recruitment and survival may contribute to the observed relationships between water availability, habitat associations and patterns of tree species richness.

Fuels and fires influence vegetation via above‐ and belowground pathways in a high‐diversity plant community

Summary Fire strongly influences plant populations and communities around the world, making it an important agent of plant evolution. Fire influences vegetation through multiple pathways, both above- and belowground. Few studies have yet attempted to tie these pathways together in a mechanistic way through soil heating even though the importance of soil heating for plants in fire-prone ecosystems is increasingly recognized. Here we combine an experimental approach with structural equation modelling (SEM) to simultaneously examine multiple pathways through which fire might influence herbaceous vegetation. In a high-diversity longleaf pine groundcover community in Louisiana, USA, we manipulated fine-fuel biomass and monitored the resulting fires with high-resolution thermocouples placed in vertical profile above- and belowground. We predicted that vegetation response to burning would be inversely related to fuel load owing to relationships among fuels, fire temperature, duration and soil heating. We found that fuel manipulations altered fire properties and vegetation responses, of which soil heating proved to be a highly accurate predictor. Fire duration acting through soil heating was important for vegetation response in our SEMs, whereas fire temperature was not. Our results indicate that in this herbaceous plant community, fire duration is a good predictor of soil heating and therefore of vegetation response to fire. Soil heating may be the key determinant of vegetation response to fire in ecosystems wherein plants persist by resprouting or reseeding from soil-stored propagules. Synthesis. Our SEMs demonstrate how the complex pathways through which fires influence plant community structure and dynamics can be examined simultaneously. Comparative studies of these pathways across different communities will provide important insights into the ecology, evolution and conservation of fire-prone ecosystems.