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      The Relative Importance of Janzen-Connell Effects in Influencing the Spatial Patterns at the Gutianshan Subtropical Forest

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          Abstract

          The Janzen-Connell hypothesis is among the most important theories put forward to explain species coexistence in species-rich communities. However, the relative importance of Janzen-Connell effects with respect to other prominent mechanisms of community assembly, such as dispersal limitation, self-thinning due to competition, or habitat association, is largely unresolved. Here we use data from a 24-ha Gutianshan subtropical forest to address it. First we tested for significant associations of adults, juveniles, and saplings with environmental variables. Second we evaluated if aggregation decreased with life stage. In a third analysis we approximately factored out the effect of habitat association and comprehensively analyzed the spatial associations of intraspecific adults and offspring (saplings, juveniles) of 46 common species at continuous neighborhood distances. We found i) that, except for one, all species were associated with at least one environmental variable during at least one of their life stages, but the frequency of significant habitat associations declined with increasing life stage; ii) a decline in aggregation with increasing life stage that was strongest from juveniles to adults; and iii) intraspecific adult-offspring associations were dominated by positive relationships at neighborhood distances up to 10 m. Our results suggest that Janzen-Connell effects were not the dominant mechanisms in structuring the spatial patterns of established trees in the subtropical Gutianshan forest. The spatial patterns may rather reflect the joint effects of size-dependent self-thinning, dispersal limitation and habitat associations. Our findings contribute to a more comprehensive understanding of the relative importance of Janzen-Connell effects in influencing plant community structure under strong topographic heterogeneity.

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          Most cited references13

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          Tree dispersion, abundance, and diversity in a tropical dry forest.

          S Hubbell (1979)
          Patterns of tree abundance and dispersion in a tropical deciduous (dry) forest are summarized. The generalization that tropical trees have spaced adults did not hold. All species were either clumped or randomly dispersed, with rare species more clumped than common species. Breeding system was unrelated to species abundance or dispersion, but clumping was related to mode of seed dispersal. Juvenile densities decreased approximately exponentially away from adults. Rare species gave evidence of poor reproductive performance compared with their performance when common in nearby forests. Patterns of relative species abundance in the dry forest are compared with patterns in other forests, and are explained by a simple stochastic model based on random-walk immigration and extinction set in motion by periodic community disturbance.
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            Pervasive density-dependent recruitment enhances seedling diversity in a tropical forest.

            Negative density-dependent recruitment of seedlings, that is, seeds of a given species are less likely to become established seedlings if the density of that species is high, has been proposed to be an important mechanism contributing to the extraordinary diversity of tropical tree communities because it can potentially prevent any particular species from usurping all available space, either in close proximity to seed sources or at relatively larger spatial scales. However, density-dependent recruitment does not necessarily enhance community diversity. Furthermore, although density-dependent effects have been found at some life stages in some species, no study has shown that density-dependent recruitment affects community diversity. Here we report the results of observations in a lowland, moist forest in the Republic of Panamá in which the species identities of 386,027 seeds that arrived at 200 seed traps were compared with the species identities of 13,068 seedlings that recruited into adjacent plots over a 4-year period. Across the 200 sites, recruit seedling diversity was significantly higher than seed diversity. Part of this difference was explained by interspecies differences in average recruitment success. Even after accounting for these differences, however, negative density-dependent recruitment contributes significantly to the increase in diversity from seeds to seedling recruits.
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              Spatial patterns in the distribution of tropical tree species.

              Fully mapped tree census plots of large area, 25 to 52 hectares, have now been completed at six different sites in tropical forests, including dry deciduous to wet evergreen forest on two continents. One of the main goals of these plots has been to evaluate spatial patterns in tropical tree populations. Here the degree of aggregation in the distribution of 1768 tree species is examined based on the average density of conspecific trees in circular neighborhoods around each tree. When all individuals larger than 1 centimeter in stem diameter were included, nearly every species was more aggregated than a random distribution. Considering only larger trees (>/= 10 centimeters in diameter), the pattern persisted, with most species being more aggregated than random. Rare species were more aggregated than common species. All six forests were very similar in all the particulars of these results.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2013
                5 September 2013
                : 8
                : 9
                : e74560
                Affiliations
                [1 ]State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing, China
                [2 ]Department of Ecological Modelling, UFZ Helmholtz Centre for Environmental Research, Leipzig, Germany
                The University of Texas at San Antonio, United States of America
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: YZ SG TW KPM. Performed the experiments: YZ HBR. Analyzed the data: YZ SG TW. Wrote the paper: YZ SG TW.

                Article
                PONE-D-12-35294
                10.1371/journal.pone.0074560
                3764046
                24040283
                77085ac5-8e8d-4ae4-b8e8-dc52567244fd
                Copyright @ 2013

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 12 November 2012
                : 2 August 2013
                Page count
                Pages: 8
                Funding
                This study was financially supported by the National Natural Science Foundation of China (31200342), Key Innovation Project of CAS (KZCX2-YW-430) and State Key Laboratory of Vegetation and Environmental Change (2011zyts12). SG was supported by the European Research Council (ERC) Advanced Grant “SpatioDiversity” (grant number 233066) to TW. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
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