Assessing Evidence for a Pervasive Alteration in Tropical Tree Communities

In Amazonian tropical forests, recent studies have reported increases in aboveground biomass and in primary productivity, as well as shifts in plant species composition favouring fast-growing species over slow-growing ones. This pervasive alteration of mature tropical forests was attributed to global environmental change, such as an increase in atmospheric CO2 concentration, nutrient deposition, temperature, drought frequency, and/or irradiance. We used standardized, repeated measurements of over 2 million trees in ten large (16–52 ha each) forest plots on three continents to evaluate the generality of these findings across tropical forests. Aboveground biomass increased at seven of our ten plots, significantly so at four plots, and showed a large decrease at a single plot. Carbon accumulation pooled across sites was significant (+0.24 MgC ha−1 y−1, 95% confidence intervals [0.07, 0.39] MgC ha−1 y−1), but lower than reported previously for Amazonia. At three sites for which we had data for multiple census intervals, we found no concerted increase in biomass gain, in conflict with the increased productivity hypothesis. Over all ten plots, the fastest-growing quartile of species gained biomass (+0.33 [0.09, 0.55] % y−1) compared with the tree community as a whole (+0.15 % y−1); however, this significant trend was due to a single plot. Biomass of slow-growing species increased significantly when calculated over all plots (+0.21 [0.02, 0.37] % y−1), and in half of our plots when calculated individually. Our results do not support the hypothesis that fast-growing species are consistently increasing in dominance in tropical tree communities. Instead, they suggest that our plots may be simultaneously recovering from past disturbances and affected by changes in resource availability. More long-term studies are necessary to clarify the contribution of global change to the functioning of tropical forests

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

Temporal population variability in local forest communities has mixed effects on tree species richness across a latitudinal gradient

Among the local processes that determine species diversity in ecological communities, fluctuation‐dependent mechanisms that are mediated by temporal variability in the abundances of species populations have received significant attention. Higher temporal variability in the abundances of species populations can increase the strength of temporal niche partitioning but can also increase the risk of species extinctions, such that the net effect on species coexistence is not clear. We quantified this temporal population variability for tree species in 21 large forest plots and found much greater variability for higher latitude plots with fewer tree species. A fitted mechanistic model showed that among the forest plots, the net effect of temporal population variability on tree species coexistence was usually negative, but sometimes positive or negligible. Therefore, our results suggest that temporal variability in the abundances of species populations has no clear negative or positive contribution to the latitudinal gradient in tree species richness

An estimate of the number of tropical tree species

The high species richness of tropical forests has long been recognized, yet there remains substantial uncertainty regarding the actual number of tropical tree species. Using a pantropical tree inventory database from closed canopy forests, consisting of 657,630 trees belonging to 11,371 species, we use a fitted value of Fisher’s alpha and an approximate pantropical stem total to estimate the minimum number of tropical forest tree species to fall between ∼40,000 and ∼53,000, i.e. at the high end of previous estimates. Contrary to common assumption, the Indo-Pacific region was found to be as species-rich as the Neotropics, with both regions having a minimum of ∼19,000–25,000 tree species. Continental Africa is relatively depauperate with a minimum of ∼4,500–6,000 tree species. Very few species are shared among the African, American, and the Indo-Pacific regions. We provide a methodological framework for estimating species richness in trees that may help refine species richness estimates of tree-dependent taxa

Global importance of large-diameter trees

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

Multidimensional tree niches in a tropical dry forest

The extent to which interspecific niche differences structure plant communities is highly debated, with extreme viewpoints ranging from fine-scaled niche partitioning, where every species in the community is specialized to a distinct niche, to neutrality, where species have no niche or fitness differences. However, there exists a default position wherein niches of species in a community are determined by their evolutionary and biogeographic histories, irrespective of other species within the community. According to this viewpoint, a broad range of pair-wise niche overlaps-from completely overlapping to completely distinct-are expected in any community without the need to invoke interspecific interactions. We develop a method that can test for both habitat associations and niche differences along an arbitrary number of spatial and temporal niche dimensions and apply it to a 24-yr data set of the eight dominant woody-plant species (representing 84% and 76% of total community abundance and basal area, respectively) from a 50-ha permanent plot in a southern Indian tropical dry forest, using edaphic, topographic, and precipitation variables as niche axes. Species separated into two broad groups in niche space-one consisting of three canopy species and the other of a canopy species and four understory species-along axes that corresponded mainly to variation in soil P, Al and a topographic index of wetness. Species within groups tended to have significantly greater niche overlap than expected by chance. Community-wide niche overlap in spatial and temporal niche axes was never smaller than expected by chance. Species-habitat associations were neither necessary nor sufficient preconditions for niche differences to be present. Our results suggest that this tropical dry-forest community consists of several tree species with broadly overlapping niches, and where significant niche differences do exist, they are not readily interpretable as evidence for niche differentiation. We argue, based on a survey of the literature, that many of the observed niche differences in tropical forests are more parsimoniously viewed as autecological differences between species that exist independently of interspecific interactions

Woody plant diversity in relation to environmental factors in a seasonally dry tropical forest landscape

QuestionsWater availability is known to be a first-order driver of plant diversity; yet water also affects fire regimes and soil fertility, which, in turn, affect plant diversity. We examined how precipitation, fire and soil properties jointly determine woody plant diversity. Specifically, we asked how woody plant diversity varies along a sharp precipitation gradient (about 600-1,800mm mean annual precipitation MAP]within a similar to 45-km distance) exhibiting considerable variation in long-term fire burn frequency and soil fertility, in a southern Indian seasonally dry tropical forest (SDTF) landscape. LocationMudumalai, Western Ghats, India. MethodsWoody plants 1-cm DBH were enumerated in 19 1-ha permanent plots spanning a range of tropical vegetation types from dry thorn forest, through dry and moist deciduous forest to semi-evergreen forest. Burn frequencies were derived from annual fire maps. Six measures of surface soil properties - total exchangeable bases (Ca+Mg+K), organic carbon (OC), total N, pH, plant available P and micronutrients (Fe+Cu+Zn+Mn) were used in the analyses. Five measures of diversity - species richness, Shannon diversity, the rarefied/extrapolated versions of these two measures, and Fisher's - were modelled as functions of MAP, annual fire burn frequency and the principal components of soil properties. ResultsMost soil nutrients and OC increased with MAP, except in the wettest sites. Woody productivity increased with MAP, while fire frequency was highest at intermediate values of MAP. Woody plant diversity increased with MAP but decreased with increasing fire frequency, resulting in two local diversity maxima along the MAP gradient - in the semi-evergreen and dry thorn forest - separated by a low-diversity central region in dry deciduous forest where fire frequency was highest. Soil variables were, on the whole, less strongly correlated with diversity than MAP. ConclusionsAlthough woody plant diversity in this landscape, representative of regional SDTFs, is primarily limited by water availability, our study emphasizes the role of fire as a potentially important second-order driver that acts to reduce diversity in this landscape

The roots of the drought: Hydrology and water uptake strategies mediate forest-wide demographic response to precipitation

ISI Document Delivery No.: GJ5TETimes Cited: 0Cited Reference Count: 60Chitra-Tarak, Rutuja Ruiz, Laurent Dattaraja, Handanakere S. Kumar, M. S. Mohan Riotte, Jean Suresh, Hebbalalu S. McMahon, Sean M. Sukumar, RamanMinistry of Environment, Forest and Climate Change, Govt. of India; CSIR, India; French Institute of Research for Development (IRD), CNRS-INSU, Toulouse University, France; NSF [1137366, 1046113]Ministry of Environment, Forest and Climate Change, Govt. of India; CSIR, India; French Institute of Research for Development (IRD), CNRS-INSU, Toulouse University, France; NSF, Grant/Award Numbers: 1137366, 1046113WileyHoboken1. Drought-induced tree mortality is expected to increase globally due to climate change, with profound implications for forest composition, function and global climate feedbacks. How drought is experienced by different species is thought to depend fundamentally on where they access water vertically below-ground, but this remains untracked so far due to the difficulty of measuring water availability at depths at which plants access water (few to several tens of metres), the broad temporal scales at which droughts at those depths unfold (seasonal to decadal), and the difficulty in linking these patterns to forest-wide species-specific demographic responses. 2. We address this problem through a new eco-hydrological framework: we used a hydrological model to estimate below-ground water availability by depth over a period of two decades that included a multi-year drought. Given this water availability scenario and 20year long-records of species-specific growth patterns, we inversely estimated the relative depths at which 12 common species in the forest accessed water via a model of water stress. Finally, we tested whether our estimates of species relative uptake depths predicted mortality in the multi-year drought. 3. The hydrological model revealed clear below-ground niches as precipitation was decoupled from water availability by depth at multi-annual scale. Species partitioned the hydrological niche by diverging in their uptake depths and so in the same forest stand, different species experienced very different drought patterns, resulting in clear differences in species-specific growth. Finally, species relative water uptake depths predicted species mortality patterns after the multi-year drought. Species that our method ranked as relying on deeper water were the ones that had suffered from greater mortality, as the zone from which they access water took longer to recharge after depletion. 4. Synthesis. This research changes our understanding of how hydrological niches operate for trees, with a trade-off between realized growth potential and survival under drought with decadal scale return time. The eco-hydrological framework highlights the importance of species-specific below-ground strategies in predicting forest response to drought. Applying this framework more broadly may help us better understand species coexistence in diverse forest communities and improve mechanistic predictions of forests productivity and compositional change under future climate

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