In many forested ecosystems, the architecture and functional ecology of certain tree species define forest structure and their species‐specific traits control ecosystem dynamics. Such foundation tree species are declining throughout the world due to introductions and outbreaks of pests and pathogens, selective removal of individual taxa, and over‐harvesting. Through a series of case studies, we show that the loss of foundation tree species changes the local environment on which a variety of other species depend; how this disrupts fundamental ecosystem processes, including rates of decomposition, nutrient fluxes, carbon sequestration, and energy flow; and dramatically alters the dynamics of associated aquatic ecosystems. Forests in which dynamics are controlled by one or a few foundation species appear to be dominated by a small number of strong interactions and may be highly susceptible to alternating between stable states following even small perturbations. The ongoing decline of many foundation species provides a set of important, albeit unfortunate, opportunities to develop the research tools, models, and metrics needed to identify foundation species, anticipate the cascade of immediate, short‐ and long‐term changes in ecosystem structure and function that will follow from their loss, and provide options for remedial conservation and management.
Phenotypic traits and their associated trade-offs have been shown to have globally consistent effects on individual plant physiological functions 1-3 , but how these effects scale up to influence competition, a key driver of community assembly in terrestrial vegetation, has remained unclear 4 . Here we use growth data from more than 3 million trees in over 140,000 plots across the world to show how three key functional traits-wood density, specific leaf area and maximum height-consistently influence competitive interactions. Fast maximum growth of a species was correlated negatively with its wood density in all biomes, and positively with its specific leaf area in most biomes. Low wood density was also correlated with a low ability to tolerate competition and a low competitive effect on neighbours, while high specific leaf area was correlated with a low competitive effect. Thus, traits generate trade-offs between performance with competition versus performance without competition, a fundamental ingredient in the classical hypothesis that the coexistence of plant species is enabled via differentiation in their successional strategies 5 . Competition within species was stronger than between species, but an increase in trait dissimilarity between species had little influence in weakening competition. No benefit of dissimilarity was detected for specific leaf area or wood density, and only a weak benefit for maximum height. Our traitbased approach to modelling competition makes generalization possible across the forest ecosystems of the world and their highly diverse species composition.Phenotypic traits are considered fundamental drivers of community assembly and thus species diversity 1,6 . The effects of traits on individual plant physiologies and functions are increasingly understood, and have been shown to be underpinned by well-known and globally consistent trade-offs 1-3 . For instance, traits such as wood density and specific leaf area capture trade-offs between the construction cost and longevity or strength of wood and leaf tissues 2,3 . By contrast, we still have a limited understanding of how such trait-based trade-offs translate into competitive interactions between species, particularly for long-lived organisms such as trees. Competition is a key filter through which ecological and evolutionary success is determined 4 . A long-standing hypothesis is that the intensity of competition decreases as two species diverge in trait values 7 (trait dissimilarity). The few studies [8][9][10][11][12][13] that have explored links between traits and competition have shown that linkages were more complex than this, as particular trait values may also confer competitive advantage independently from trait dissimilarity 9,13,14 . This distinction is fundamental for species coexistence and the local mixture of traits. If neighbourhood competition is driven mainly by trait dissimilarity, this will favour a wide spread of trait values at a local scale. By contrast, if neighbourhood interactions are mainly driven by the c...
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 oldgrowth 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.
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 AE30% change), atmospheric deposition of nitrogen and sulfur compounds (up to 3.8 g N m À2 yr À1 and 3.1 g S m À2 yr À1), 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 CTFSForestGEO 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.
The problem of scale dependency is widespread in investigations of ecological communities. Null model investigations of community assembly exemplify the challenges involved because they typically include subjectively defined "regional species pools." The burgeoning field of community phylogenetics appears poised to face similar challenges. Our objective is to quantify the scope of the problem of scale dependency by comparing the phylogenetic structure of assemblages across contrasting geographic and taxonomic scales. We conduct phylogenetic analyses on communities within three tropical forests, and perform a sensitivity analysis with respect to two scaleable inputs: taxonomy and species pool size. We show that (1) estimates of phylogenetic overdispersion within local assemblages depend strongly on the taxonomic makeup of the local assemblage and (2) comparing the phylogenetic structure of a local assemblage to a species pool drawn from increasingly larger geographic scales results in an increased signal of phylogenetic clustering. We argue that, rather than posing a problem, "scale sensitivities" are likely to reveal general patterns of diversity that could help identify critical scales at which local or regional influences gain primacy for the structuring of communities. In this way, community phylogenetics promises to fill an important gap in community ecology and biogeography research.
Aim Tropical forests store 25% of global carbon and harbour 96% of the world's tree species, but it is not clear whether this high biodiversity matters for carbon storage. Few studies have teased apart the relative importance of forest attributes and environmental drivers for ecosystem functioning, and no such study exists for the tropics. Location Neotropics. Methods We relate aboveground biomass (AGB) to forest attributes (diversity and structure) and environmental drivers (annual rainfall and soil fertility) using data from 144,000 trees, 2050 forest plots and 59 forest sites. The sites span the complete latitudinal and climatic gradients in the lowland Neotropics, with rainfall ranging from 750 to 4350 mm year−1. Relationships were analysed within forest sites at scales of 0.1 and 1 ha and across forest sites along large‐scale environmental gradients. We used a structural equation model to test the hypothesis that species richness, forest structural attributes and environmental drivers have independent, positive effects on AGB. Results Across sites, AGB was most strongly driven by rainfall, followed by average tree stem diameter and rarefied species richness, which all had positive effects on AGB. Our indicator of soil fertility (cation exchange capacity) had a negligible effect on AGB, perhaps because we used a global soil database. Taxonomic forest attributes (i.e. species richness, rarefied richness and Shannon diversity) had the strongest relationships with AGB at small spatial scales, where an additional species can still make a difference in terms of niche complementarity, while structural forest attributes (i.e. tree density and tree size) had strong relationships with AGB at all spatial scales. Main conclusions Biodiversity has an independent, positive effect on AGB and ecosystem functioning, not only in relatively simple temperate systems but also in structurally complex hyperdiverse tropical forests. Biodiversity conservation should therefore be a key component of the UN Reducing Emissions from Deforestation and Degradation strategy.
Abstract. We present a likelihood-based regression method that was developed to analyze the effects of neighborhood competitive interactions and hurricane damage on tree growth and survival. The purpose of the method is to provide robust parameter estimates for a spatially explicit forest simulator and to gain insight into the processes that drive the patterns of species abundance in tropical forests. We test the method using census data from the 16-ha Luquillo Forest Dynamics Plot in Puerto Rico and describe effects of the spatial configuration, sizes, and species of neighboring trees on the growth and survival of 12 dominant tree species representing a variety of life history strategies. Variation in sizedependent growth and mortality suggests a complex relationship between size, growth, and survival under different regimes of light availability. Crowding effects on growth and survival appear to be idiosyncratic to each individual species, and with the exception of pioneers, there is little commonality among species that share similar life histories.We also explain the implications of differential susceptibility to hurricane damage on species' growth and survival and on their ability to respond to damage to neighboring trees. Tree species in the Luquillo Forest Dynamics Plot differ strikingly in both their susceptibility to hurricane disturbance and the nature of their recovery from wind disturbance, through response of both adult plants and juveniles to enhanced resource availability. At the stand level, intense competitive thinning of densely packed saplings that grew after hurricane damage accounted for the majority of post-hurricane mortality, particularly for shadeintolerant species. At the individual species level, effects of previous hurricane damage on growth and survival depended primarily on variation in the quantity and quality of hurricane damage sustained by target species and their interaction with life history characteristics of these individual species.Finally, we compare models that make different assumptions about the effects of competing species on tree growth and survival (e.g., equivalence of competitors vs. distinct species-specific effects). Size effects alone could not account for growth and survival for the majority of target species. Our results also demonstrate that competing species have distinct per capita effects on growth of dominant target species. In contrast, we found moderate support for a model that assumed functional equivalence of competitors on survival.
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