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Stephen P Hubbell

Publications and source records attributed to Stephen P Hubbell.

14 recordsLinked to original sources

Soil nutrients influence spatial distributions of tropical tree species.

The importance of niche vs. neutral assembly mechanisms in structuring tropical tree communities remains an important unsettled question in community ecology [Bell G (2005) Ecology 86:1757-1770]. There is ample evidence that species distributions are determined by soils and habitat factors at landscape (<10(4) km(2)) and regional scales. At local scales (<1 km(2)), however, habitat factors and species distributions show comparable spatial aggregation, making it difficult to disentangle the importance of niche and dispersal processes. In this article, we test soil resource-based niche assembly at a local scale, using species and soil nutrient distributions obtained at high spatial resolution in three diverse neotropical forest plots in Colombia (La Planada), Ecuador (Yasuni), and Panama (Barro Colorado Island). Using spatial distribution maps of >0.5 million individual trees of 1,400 species and 10 essential plant nutrients, we used Monte Carlo simulations of species distributions to test plant-soil associations against null expectations based on dispersal assembly. We found that the spatial distributions of 36-51% of tree species at these sites show strong associations to soil nutrient distributions. Neutral dispersal assembly cannot account for these plant-soil associations or the observed niche breadths of these species. These results indicate that belowground resource availability plays an important role in the assembly of tropical tree communities at local scales and provide the basis for future investigations on the mechanisms of resource competition among tropical tree species.

Ecosystem↗

Genetic evidence of frequent long-distance recruitment in a vertebrate-dispersed tree.

The importance of dispersal for the maintenance of biodiversity, while long-recognized, has remained unresolved. We used molecular markers to measure effective dispersal in a natural population of the vertebrate-dispersed Neotropical tree, Simarouba amara (Simaroubaceae) by comparing the distances between maternal parents and their offspring and comparing gene movement via seed and pollen in the 50 ha plot of the Barro Colorado Island forest, Central Panama. In all cases (parent-pair, mother-offspring, father-offspring, sib-sib) distances between related pairs were significantly greater than distances to nearest possible neighbours within each category. Long-distance seedling establishment was frequent: 74% of assigned seedlings established > 100 m from the maternal parent [mean = 392 +/- 234.6 m (SD), range = 9.3-1000.5 m] and pollen-mediated gene flow was comparable to that of seed [mean = 345.0 +/- 157.7 m (SD), range 57.6-739.7 m]. For S. amara we found approximately a 10-fold difference between distances estimated by inverse modelling and mean seedling recruitment distances (39 m vs. 392 m). Our findings have important implications for future studies in forest demography and regeneration, with most seedlings establishing at distances far exceeding those demonstrated by negative density-dependent effects.

Animals↗

Testing metabolic ecology theory for allometric scaling of tree size, growth and mortality in tropical forests.

The theory of metabolic ecology predicts specific relationships among tree stem diameter, biomass, height, growth and mortality. As demographic rates are important to estimates of carbon fluxes in forests, this theory might offer important insights into the global carbon budget, and deserves careful assessment. We assembled data from 10 old-growth tropical forests encompassing censuses of 367 ha and > 1.7 million trees to test the theory's predictions. We also developed a set of alternative predictions that retained some assumptions of metabolic ecology while also considering how availability of a key limiting resource, light, changes with tree size. Our results show that there are no universal scaling relationships of growth or mortality with size among trees in tropical forests. Observed patterns were consistent with our alternative model in the one site where we had the data necessary to evaluate it, and were inconsistent with the predictions of metabolic ecology in all forests.

Biometry↗

Comparing tropical forest tree size distributions with the predictions of metabolic ecology and equilibrium models.

Tropical forests vary substantially in the densities of trees of different sizes and thus in above-ground biomass and carbon stores. However, these tree size distributions show fundamental similarities suggestive of underlying general principles. The theory of metabolic ecology predicts that tree abundances will scale as the -2 power of diameter. Demographic equilibrium theory explains tree abundances in terms of the scaling of growth and mortality. We use demographic equilibrium theory to derive analytic predictions for tree size distributions corresponding to different growth and mortality functions. We test both sets of predictions using data from 14 large-scale tropical forest plots encompassing censuses of 473 ha and > 2 million trees. The data are uniformly inconsistent with the predictions of metabolic ecology. In most forests, size distributions are much closer to the predictions of demographic equilibrium, and thus, intersite variation in size distributions is explained partly by intersite variation in growth and mortality.

Biomass↗

Neutral theory and the evolution of ecological equivalence.

Since the publication of the unified neutral theory in 2001, there has been much discussion of the theory, pro and con. The hypothesis of ecological equivalence is the fundamental yet controversial idea behind neutral theory. Assuming trophically similar species are demographically alike (symmetric) on a per capita basis is only an approximation, but it is equivalent to asking: How many of the patterns of ecological communities are the result of species similarities, rather than of species differences? The strategy behind neutral theory is to see how far one can get with the simplification of assuming ecological equivalence before introducing more complexity. In another paper, I review the empirical evidence that led me to hypothesize ecological equivalence among many of the tree species in the species-rich tropical forest on Barro Colorado Island (BCI). In this paper, I develop a simple model for the evolution of ecological equivalence or niche convergence, using as an example evolution of the suite of life history traits characteristic of shade tolerant tropical tree species. Although the model is simple, the conclusions from it seem likely to be robust. I conclude that ecological equivalence for resource use are likely to evolve easily and often, especially in species-rich communities that are dispersal and recruitment limited. In the case of the BCI forest, tree species are strongly dispersal- and recruitment-limited, not only because of restricted seed dispersal, but also because of low recruitment success due to heavy losses of the seedling stages to predators and pathogens and other abiotic stresses such as drought. These factors and the high species richness of the community strongly reduce the potential for competitive exclusion of functionally equivalent or nearly equivalent species.

Biodiversity↗

Contrasting structure and composition of the understory in species-rich tropical rain forests.

In large samples of trees > or = 1 cm dbh (more than 1 million trees and 3000 species), in six lowland tropical forests on three continents, we assigned species with >30 individuals to one of six classes of stature at maturity (SAM). We then compared the proportional representation of understory trees (1-2 cm dbh) among these classes. The understory of the three Asian sites was predominantly composed of the saplings of large-canopy trees whereas the African and American sites were more richly stocked with trees of the smaller SAM classes. Differences in class representation were related to taxonomic families that were present exclusively in one continent or another. Families found in the Asian plots but not in the American plot (e.g., Dipterocarpaceae, Fagaceae) were predominantly species of the largest SAM classes, whereas families exclusive to the American plots (e.g., Melastomataceae sensu stricto, Piperaceae, and Malvaceae [Bombacacoidea]) were predominantly species of small classes. The African plot was similar to Asia in the absence of those American families rich in understory species, while similar to America in lacking the Asian families rich in canopy species. The numerous understory species of Africa were chiefly derived from families shared with Asia and/or America. The ratio of saplings (1-2 cm dbh) to conspecific canopy trees (>40 cm dbh) was lower in American plots than in the Asian plots. Possible explanations for these differences include phenology, moisture and soil fertility regimes, phyletic constraints, and the role of early successional plants in forest development. These results demonstrate that tropical forests that appear similar in tree number, basal area, and the family taxonomy of canopy trees nonetheless differ in ecological structure in ways that may impact the ecology of pollinators, dispersers, and herbivores and might reflect fundamental differences in canopy tree regeneration.

Biodiversity↗

The phylogenetic structure of a neotropical forest tree community.

Numerous ecological and evolutionary processes are thought to play a role in maintaining the high plant species diversity of tropical forests. An understanding of the phylogenetic structure of an ecological community can provide insights into the relative importance of different processes structuring that community. The objectives of this study were to measure the phylogenetic structure of Neotropical forest tree communities in the Forest Dynamics Plot (FDP) on Barro Colorado Island, Panama, to determine how the phylogenetic structure of tree communities varied among spatial scales and habitats within the FDP, and to study the effects of null-model choice on estimates of community phylogenetic structure. We measured community phylogenetic structure for tree species occurring together in quadrats ranging in size from 10 x 10 m to 100 X 100 m in the FDP. We estimated phylogenetic structure by comparing observed phylogenetic distances among species to the distribution of phylogenetic distances for null communities generated using two different null models. A null model that did not maintain observed species occurrence frequencies tended to find nonrandom community phylogenetic structure, even for random data. Using a null model that maintained observed species frequencies in null communities, the average phylogenetic structure of tree communities in the FDP was close to random at all spatial scales examined, but more quadrats than expected contained species that were phylogenetically clustered or overdispersed, and phylogenetic structure varied among habitats. In young forests and plateau habitats, communities were phylogenetically clustered, meaning that trees were more closely related to their neighbors than expected, while communities in swamp and slope habitats were phylogenetically overdispersed, meaning that trees were more distantly related to their neighbors than expected. Phylogenetic clustering suggests the importance of environmental filtering of phylogenetically conserved traits in young forests and plateau habitats, but the phylogenetic overdispersion observed in other habitats has several possible explanations, including variation in the strength of ecological processes among habitats or the phylogenetic history of niches, traits, and habitat associations. Future studies will need to include information on species traits in order to explain the variation in phylogenetic structure among habitats in tropical forests.

Ecosystem↗

Density dependence explains tree species abundance and diversity in tropical forests.

The recurrent patterns in the commonness and rarity of species in ecological communities--the relative species abundance--have puzzled ecologists for more than half a century. Here we show that the framework of the current neutral theory in ecology can easily be generalized to incorporate symmetric density dependence. We can calculate precisely the strength of the rare-species advantage that is needed to explain a given RSA distribution. Previously, we demonstrated that a mechanism of dispersal limitation also fits RSA data well. Here we compare fits of the dispersal and density-dependence mechanisms for empirical RSA data on tree species in six New and Old World tropical forests and show that both mechanisms offer sufficient and independent explanations. We suggest that RSA data cannot by themselves be used to discriminate among these explanations of RSA patterns--empirical studies will be required to determine whether RSA patterns are due to one or the other mechanism, or to some combination of both.

Biodiversity↗

Spatial scaling in model plant communities.

We present an analytically tractable variant of the voter model that provides a quantitatively accurate description of Beta diversity (two-point correlation function) in two tropical forests. The model exhibits novel scaling behavior that leads to links between ecological measures such as relative species abundance and the species-area relationship.

Biodiversity↗

Neutral theory: the stability of forest biodiversity.

The unified neutral theory of biodiversity and biogeography provides a dynamic null hypothesis for the assembly of natural communities. It is also useful for understanding the influence of speciation, extinction, dispersal and ecological drift on patterns of relative species abundance, species-area relationships and phylogeny. Clark and McLachlan argue that neutral drift is inconsistent with the palaeorecord of stability in fossil pollen assemblages of the Holocene forests of southern Canada. We show here that their analysis is based on a partial misunderstanding of neutral theory and that their data alone cannot unambiguously test its validity.

Biodiversity↗

Estimating local interaction from spatiotemporal forest data, and Monte Carlo bias correction.

We point out a general problem in fitting continuous time spatially explicit models to a temporal sequence of spatial data observed at discrete times. To illustrate the problem, we examined the continuous time Markov model for forest gap dynamics. A forest is assumed to be apportioned into discrete cells (or sites) arranged in a regular square lattice. Each site is characterized as either a gap or a non-gap site according to the vegetation height of trees. The model incorporates the influence of neighboring sites on transition rate: transition rate from a non-gap to a gap site increases linearly with the number of neighbors that are currently in the gap state, and vice versa. We fitted the model to the spatiotemporal data of canopy height observed at the permanent plot in Barro Colorado Island (BCI). When we used the approximate maximum likelihood method to estimate the parameters of the model, the estimated transition rates included a large bias-in particular, the strength of interaction between nearby sites was underestimated. This bias originated from the assumption that each transition between two observation times is independent. The interaction between sites at local scale creates a long chain of transitions within a single census interval, which violates the independence of each transition. We show that a computer-intensive method, called Monte Carlo bias correction (MCBC), is very effective in removing the bias included in the estimate. The global and local gap densities measuring spatial aggregation of gap sites were computed from simulated and real gap dynamics to assess the model. When the approximate likelihood estimates were applied to the model, the predicted local gap density was clearly lower than the observed one. The use of MCBC estimates, suggesting a strong interaction between sites, improved this discrepancy.

Data Interpretation, Statistical↗

Percolation theory for the distribution and abundance of species.

We develop and test new models that unify the mathematical relationships among the abundance of a species, the spatial dispersion of the species, the number of patches occupied by the species, the edge length of the occupied patches, and the scale on which the distribution of species is mapped. The models predict that species distributions will exhibit percolation critical thresholds, i.e., critical population abundances at which the fragmented patches (as measured by the number of patches and edge length) start to coalesce to form large patches.

Croton↗

Neutral theory and relative species abundance in ecology.

The theory of island biogeography asserts that an island or a local community approaches an equilibrium species richness as a result of the interplay between the immigration of species from the much larger metacommunity source area and local extinction of species on the island (local community). Hubbell generalized this neutral theory to explore the expected steady-state distribution of relative species abundance (RSA) in the local community under restricted immigration. Here we present a theoretical framework for the unified neutral theory of biodiversity and an analytical solution for the distribution of the RSA both in the metacommunity (Fisher's log series) and in the local community, where there are fewer rare species. Rare species are more extinction-prone, and once they go locally extinct, they take longer to re-immigrate than do common species. Contrary to recent assertions, we show that the analytical solution provides a better fit, with fewer free parameters, to the RSA distribution of tree species on Barro Colorado Island, Panama, than the lognormal distribution.

Animals↗

Beta-diversity in tropical forest trees.

The high alpha-diversity of tropical forests has been amply documented, but beta-diversity-how species composition changes with distance-has seldom been studied. We present quantitative estimates of beta-diversity for tropical trees by comparing species composition of plots in lowland terra firme forest in Panama, Ecuador, and Peru. We compare observations with predictions derived from a neutral model in which habitat is uniform and only dispersal and speciation influence species turnover. We find that beta-diversity is higher in Panama than in western Amazonia and that patterns in both areas are inconsistent with the neutral model. In Panama, habitat variation appears to increase species turnover relative to Amazonia, where unexpectedly low turnover over great distances suggests that population densities of some species are bounded by as yet unidentified processes. At intermediate scales in both regions, observations can be matched by theory, suggesting that dispersal limitation, with speciation, influences species turnover.

Ecosystem↗