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 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.

Tropical forests are global centres of biodiversity and carbon storage. Many tropical countries aspire to protect forest to fulfil biodiversity and climate mitigation policy targets, but the conservation strategies needed to achieve these two functions depend critically on the tropical forest tree diversity-carbon storage relationship. Assessing this relationship is challenging due to the scarcity of inventories where carbon stocks in aboveground biomass and species identifications have been simultaneously and robustly quantified. Here, we compile a unique pan-tropical dataset of 360 plots located in structurally intact old-growth closed-canopy forest, surveyed using standardised methods, allowing a multi-scale evaluation of diversity-carbon relationships in tropical forests. Diversity-carbon relationships among all plots at 1 ha scale across the tropics are absent, and within continents are either weak (Asia) or absent (Amazonia, Africa). A weak positive relationship is detectable within 1 ha plots, indicating that diversity effects in tropical forests may be scale dependent. The absence of clear diversity-carbon relationships at scales relevant to conservation planning means that carbon-centred conservation strategies will inevitably miss many high diversity ecosystems. As tropical forests can have any combination of tree diversity and carbon stocks both require explicit consideration when optimising policies to manage tropical carbon and biodiversity.

Aim To examine the contribution of large‐diameter trees to biomass, stand structure, and species richness across forest biomes. Location Global. Time period Early 21st century. Major taxa studied Woody plants. Methods We examined the contribution of large trees to forest density, richness and biomass using a global network of 48 large (from 2 to 60 ha) forest plots representing 5,601,473 stems across 9,298 species and 210 plant families. This contribution was assessed using three metrics: the largest 1% of trees ≥ 1 cm diameter at breast height (DBH), all trees ≥ 60 cm DBH, and those rank‐ordered largest trees that cumulatively comprise 50% of forest biomass. Results Averaged across these 48 forest plots, the largest 1% of trees ≥ 1 cm DBH comprised 50% of aboveground live biomass, with hectare‐scale standard deviation of 26%. Trees ≥ 60 cm DBH comprised 41% of aboveground live tree biomass. The size of the largest trees correlated with total forest biomass (r2 = .62, p < .001). Large‐diameter trees in high biomass forests represented far fewer species relative to overall forest richness (r2 = .45, p < .001). Forests with more diverse large‐diameter tree communities were comprised of smaller trees (r2 = .33, p < .001). Lower large‐diameter richness was associated with large‐diameter trees being individuals of more common species (r2 = .17, p = .002). The concentration of biomass in the largest 1% of trees declined with increasing absolute latitude (r2 = .46, p < .001), as did forest density (r2 = .31, p < .001). Forest structural complexity increased with increasing absolute latitude (r2 = .26, p < .001). Main conclusions Because large‐diameter trees constitute roughly half of the mature forest biomass worldwide, their dynamics and sensitivities to environmental change represent potentially large controls on global forest carbon cycling. We recommend managing forests for conservation of existing large‐diameter trees or those that can soon reach large diameters as a simple way to conserve and potentially enhance ecosystem services.

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.

Most ecological hypotheses about species coexistence hinge on species differences, but quantifying trait differences across species in diverse communities is often unfeasible. We examined the variation of demographic traits using a global tropical forest data set covering 4500 species in 10 large-scale tree inventories. With a hierarchical Bayesian approach, we quantified the distribution of mortality and growth rates of all tree species at each site. This allowed us to test the prediction that demographic differences facilitate species richness, as suggested by the theory that a tradeoff between high growth and high survival allows species to coexist. Contrary to the prediction, the most diverse forests had the least demographic variation. Although demographic differences may foster coexistence, they do not explain any of the 16-fold variation in tree species richness observed across the tropics. C omparative studies of tree demography typically consider the entire community as a unit, ignoring species differences (1), simply because most tree inventories include small samples of many species (2, 3). Comparative studies show that tropical forests typically have higher turnover than do temperate forests (4) and that higher tree turnover associates with higher tree diversity (5). These studies cannot, however, test ecological hypotheses about diversity, coexistence, and demography (6-10).A tradeoff between rapid growth and long life span permits species coexistence and can foster diversity: Species reproducing early in life persist despite poor competitive ability by growing rapidly on disturbed sites where resources are abundant. Long-lived species coexist by outliving the weedy invaders, persisting where resources are scarce. This is a familiar and widely known tradeoff in plant and animal communities (9-11) called the successionalniche hypothesis (7,12). At a deterministic equilibrium, an indefinite number of species can coexist by this mechanism, each differing from all others along a continuum from short life span (with high growth) to long life span (and low growth). With stochastic demography, however, there is limiting similarity and the equilibrium species richness is finite (11, 13). This hypothesis is widely quoted as an explanation for tropical forest diversity (14-16). Here, we ask whether species differences along a demographic axis explain why some tropical forests have many more species than others.If demographic niches are a key force controlling forest diversity, then more diverse forests have more demographic niches. More niches could come about either by spreading demographic rates over a wider range or packing more in the same range. Here, we focus on the first prediction: Tropical forests gain diversity by having a wider range of demographic niches, as reflected by the range of mortality and growth rates across species.We provide a direct test by quantifying mortality and growth of 4500 tree species in 10 different forests in America, Asia, and Africa (17). The 10 sites form a large-scale ob...

A 52 hectare permanent research plot was established in Lambir Hills National Park to enable long-term study of factors controlling the origin and maintenance of tree diversity. In this chapter we summarize some of our recent work on the relationships between floristic variation and edaphic heterogeneity in the Lambir forest. First, we provide a general description of the floristic composition of this hyperdiverse forest. Second, we use a detailed survey of soil chemistry to test whether floristic composition changes in relation to edaphic characteristics. We also assess the extent to which individual species have non-random distributions in the forest with respect to edaphic heterogeneity. Finally, to investigate the influence of habitat variation on floristic diversity we compare our results from the heterogeneous forest at Lambir Hills with a more homogeneous forest in Peninsular Malaysia.

Long-term surveys of entire communities of species are needed to measure fluctuations in natural populations and elucidate the mechanisms driving population dynamics and community assembly. We analysed changes in abundance of over 4000 tree species in 12 forests across the world over periods of 6-28 years. Abundance fluctuations in all forests are large and consistent with population dynamics models in which temporal environmental variance plays a central role. At some sites we identify clear environmental drivers, such as fire and drought, that could underlie these patterns, but at other sites there is a need for further research to identify drivers. In addition, cross-site comparisons showed that abundance fluctuations were smaller at species-rich sites, consistent with the idea that stable environmental conditions promote higher diversity. Much community ecology theory emphasises demographic variance and niche stabilisation; we encourage the development of theory in which temporal environmental variance plays a central role.