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      Ecological Importance of Small-Diameter Trees to the Structure, Diversity and Biomass of a Tropical Evergreen Forest at Rabi, Gabon

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          Abstract

          Tropical forests have long been recognized for their biodiversity and ecosystem services. Despite their importance, tropical forests, and particularly those of central Africa, remain understudied. Until recently, most forest inventories in Central Africa have focused on trees ≥10 cm in diameter, even though several studies have shown that small-diameter tree population may be important to demographic rates and nutrient cycling. To determine the ecological importance of small-diameter trees in central African forests, we used data from a 25-ha permanent plot that we established in the rainforest of Gabon to study the diversity and dynamics of these forests. Within the plot, we censused 175,830 trees ≥1 cm dbh from 54 families, 192 genera, and 345 species. Average tree density was 7,026 trees/ha, basal area 31.64 m 2/ha, and above-ground biomass 369.40 Mg/ha. Fabaceae, Ebenaceae and Euphorbiaceae were the most important families by basal area, density and above-ground biomass. Small-diameter trees (1 cm ≥ dbh <10 cm) comprised 93.7% of the total tree population, 16.5% of basal area, and 4.8% of the above-ground biomass. They also had diversity 18% higher at family level, 34% higher at genus level, and 42% higher at species level than trees ≥10 cm dbh. Although the relative contribution of small-diameter trees to biomass was comparable to other forests globally, their contribution to forest density, and diversity was disproportionately higher. The high levels of diversity within small-diameter classes may give these forests high levels of structural resilience to anthropogenic/natural disturbance and a changing climate.

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          Functional traits and the growth-mortality trade-off in tropical trees.

          A trade-off between growth and mortality rates characterizes tree species in closed canopy forests. This trade-off is maintained by inherent differences among species and spatial variation in light availability caused by canopy-opening disturbances. We evaluated conditions under which the trade-off is expressed and relationships with four key functional traits for 103 tree species from Barro Colorado Island, Panama. The trade-off is strongest for saplings for growth rates of the fastest growing individuals and mortality rates of the slowest growing individuals (r2 = 0.69), intermediate for saplings for average growth rates and overall mortality rates (r2 = 0.46), and much weaker for large trees (r2 80% of the explained variation and, after WD was included, LMA and H(max) made insignificant contributions. Virtually the full range of values of SM, LMA, and H(max) occurred at all positions on the growth-mortality trade-off. Although WD provides a promising start, a successful trait-based ecology of tropical forest trees will require consideration of additional traits.
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            CTFS-ForestGEO: a worldwide network monitoring forests in an era of global change.

            Global change is impacting forests worldwide, threatening biodiversity and ecosystem services including climate regulation. Understanding how forests respond is critical to forest conservation and climate protection. This review describes an international network of 59 long-term forest dynamics research sites (CTFS-ForestGEO) useful for characterizing forest responses to global change. Within very large plots (median size 25 ha), all stems ≥ 1 cm diameter are identified to species, mapped, and regularly recensused according to standardized protocols. CTFS-ForestGEO spans 25 °S-61 °N latitude, is generally representative of the range of bioclimatic, edaphic, and topographic conditions experienced by forests worldwide, and is the only forest monitoring network that applies a standardized protocol to each of the world's major forest biomes. Supplementary standardized measurements at subsets of the sites provide additional information on plants, animals, and ecosystem and environmental variables. CTFS-ForestGEO sites are experiencing multifaceted anthropogenic global change pressures including warming (average 0.61 °C), changes in precipitation (up to ± 30% change), atmospheric deposition of nitrogen and sulfur compounds (up to 3.8 g N m(-2) yr(-1) and 3.1 g S m(-2) yr(-1)), and forest fragmentation in the surrounding landscape (up to 88% reduced tree cover within 5 km). The broad suite of measurements made at CTFS-ForestGEO sites makes it possible to investigate the complex ways in which global change is impacting forest dynamics. Ongoing research across the CTFS-ForestGEO network is yielding insights into how and why the forests are changing, and continued monitoring will provide vital contributions to understanding worldwide forest diversity and dynamics in an era of global change.
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              Ecological Importance of Large-Diameter Trees in a Temperate Mixed-Conifer Forest

              Large-diameter trees dominate the structure, dynamics and function of many temperate and tropical forests. Although both scaling theory and competition theory make predictions about the relative composition and spatial patterns of large-diameter trees compared to smaller diameter trees, these predictions are rarely tested. We established a 25.6 ha permanent plot within which we tagged and mapped all trees ≥1 cm dbh, all snags ≥10 cm dbh, and all shrub patches ≥2 m2. We sampled downed woody debris, litter, and duff with line intercept transects. Aboveground live biomass of the 23 woody species was 507.9 Mg/ha, of which 503.8 Mg/ha was trees (SD = 114.3 Mg/ha) and 4.1 Mg/ha was shrubs. Aboveground live and dead biomass was 652.0 Mg/ha. Large-diameter trees comprised 1.4% of individuals but 49.4% of biomass, with biomass dominated by Abies concolor and Pinus lambertiana (93.0% of tree biomass). The large-diameter component dominated the biomass of snags (59.5%) and contributed significantly to that of woody debris (36.6%). Traditional scaling theory was not a good model for either the relationship between tree radii and tree abundance or tree biomass. Spatial patterning of large-diameter trees of the three most abundant species differed from that of small-diameter conspecifics. For A. concolor and P. lambertiana, as well as all trees pooled, large-diameter and small-diameter trees were spatially segregated through inter-tree distances <10 m. Competition alone was insufficient to explain the spatial patterns of large-diameter trees and spatial relationships between large-diameter and small-diameter trees. Long-term observations may reveal regulation of forest biomass and spatial structure by fire, wind, pathogens, and insects in Sierra Nevada mixed-conifer forests. Sustaining ecosystem functions such as carbon storage or provision of specialist species habitat will likely require different management strategies when the functions are performed primarily by a few large trees as opposed to many smaller trees.
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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, CA USA )
                1932-6203
                17 May 2016
                2016
                : 11
                : 5
                : e0154988
                Affiliations
                [1 ]Institut de Recherche en Écologie Tropicale (IRET), Libreville, Gabon
                [2 ]Wildland Resources Department, Utah State University, Logan, Utah, United States of America
                [3 ]Center for Conservation and Sustainability, Smithsonian Conservation Biology Institute, Gamba, Gabon
                [4 ]Center for Conservation and Sustainability, Smithsonian Conservation Biology Institute, Washington DC, United States of America
                [5 ]Center for Tropical Forest Science -Forest Global Earth Observatory, Smithsonian Tropical Research Institute, Smithsonian Institution, Washington DC, United States of America
                Chinese Academy of Forestry, CHINA
                Author notes

                Competing Interests: Shell Gabon provided financial support for the work. There are no patents, products in development or marketed products to declare. This does not alter the authors' adherence to PLOS ONE policies on sharing data and materials.

                Conceived and designed the experiments: AA DK HRM LK. Performed the experiments: HRM DK AA LK. Analyzed the data: DK HRM JAL. Contributed reagents/materials/analysis tools: HRM DK JAL. Wrote the paper: DK HRM JAL.

                Article
                PONE-D-16-01082
                10.1371/journal.pone.0154988
                4871370
                27186658
                65fbc499-cf99-40ce-af26-1ae932b0f240

                This is an open access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication.

                History
                : 14 January 2016
                : 22 April 2016
                Page count
                Figures: 5, Tables: 5, Pages: 15
                Funding
                Funded by: Shell (GA)
                Funded by: CTFS-ForestGEO
                Shell Gabon provided financial support for field work, especially to HRM and LK. Shell Gabon also provided logistical support including flights, lodging and food to LK, HRM, DK and 14 other field staff. Shell Gabon had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. The Smithsonian Tropical Research institute through the Forest Global Earth Observatory (ForestGEO) provided funding for the salaries of 12 field staff including HRM, and also provided funding to DK for training and technical advice. The Rabi plot was established with the protocols developed by ForestGEO, and DK is African Program Coordinator for ForestGEO. The data was analyzed by HRM, DK and JL during a series of analytical workshops funded by ForestGEO in China, US and Panama.
                Categories
                Research Article
                Biology and Life Sciences
                Organisms
                Plants
                Trees
                Biology and Life Sciences
                Ecology
                Ecosystems
                Forests
                Ecology and Environmental Sciences
                Ecology
                Ecosystems
                Forests
                Ecology and Environmental Sciences
                Terrestrial Environments
                Forests
                Biology and Life Sciences
                Ecology
                Ecological Metrics
                Biomass (Ecology)
                Ecology and Environmental Sciences
                Ecology
                Ecological Metrics
                Biomass (Ecology)
                Biology and Life Sciences
                Ecology
                Forest Ecology
                Ecology and Environmental Sciences
                Ecology
                Forest Ecology
                Biology and Life Sciences
                Plant Science
                Dendrology
                People and Places
                Geographical Locations
                Africa
                Gabon
                Biology and Life Sciences
                Ecology
                Biodiversity
                Ecology and Environmental Sciences
                Ecology
                Biodiversity
                Biology and Life Sciences
                Ecology
                Ecosystems
                Forests
                Temperate Forests
                Ecology and Environmental Sciences
                Ecology
                Ecosystems
                Forests
                Temperate Forests
                Ecology and Environmental Sciences
                Terrestrial Environments
                Forests
                Temperate Forests
                Ecology and Environmental Sciences
                Terrestrial Environments
                Temperate Forests
                Custom metadata
                The Rabi 25-hectare Forest Monitoring Plot data belongs to the plot PIs and is managed by the CTFS-ForestGEO network. Data are available under request at ( http://www.ctfs.si.edu/site/Rabi).

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