Background

The analysis of tree-ring carbon isotope composition (δ(13)C) has been widely used to estimate spatio-temporal variations in intrinsic water use efficiency (iWUE) of tree species. Numerous studies have reported widespread increases in iWUE coinciding with rising atmospheric CO(2) over the past century. While this could represent a coherent global response, the fact that increases of similar magnitude were observed across biomes with no apparent effect on tree growth raises the question of whether iWUE calculations reflect actual physiological responses to elevated CO(2) levels.

Methodology/results

Here we use Monte Carlo simulations to test if an artifact of calculation could explain observed increases in iWUE. We show that highly significant positive relationships between iWUE and CO(2) occur even when simulated data (randomized δ(13)C values spanning the observed range) are used in place of actual tree-ring δ(13)C measurements. From simulated data sets we calculated non-physiological changes in iWUE from 1900 to present and across a 4000 m altitudinal range. This generated results strikingly similar to those reported in recent studies encompassing 22 species from tropical, subtropical, temperate, boreal and mediterranean ecosystems. Only 6 of 49 surveyed case studies showed increases in iWUE significantly higher than predicted from random values.

Conclusions/significance

Our results reveal that increases in iWUE estimated from tree-ring δ(13)C occur independently of changes in (13)C discrimination that characterize physiological responses to elevated CO(2). Due to a correlation with CO(2) concentration, which is used as an independent factor in the iWUE calculation, any tree-ring δ(13)C data set would inevitably generate increasing iWUE over time. Therefore, although consistent, previously reported trends in iWUE do not necessarily reflect a coherent global response to rising atmospheric CO(2). We discuss the significance of these findings and suggest ways to distinguish real from artificial responses in future studies.

The Hawaiian Islands are an ideal location to study the response of tropical forests to climate variability because of their extreme isolation in the middle of the Pacific, which makes them especially sensitive to El Niño-Southern Oscillation (ENSO). Most research examining the response of tropical forests to drought or El Niño have focused on rainforests, however, tropical dry forests cover a large area of the tropics and may respond very differently than rainforests. We use satellite-derived Normalized Difference Vegetation Index (NDVI) from February 2000-February 2009 to show that rainforests and dry forests in the Hawaiian Islands exhibit asynchronous responses in leaf phenology to seasonal and El Niño-driven drought. Dry forest NDVI was more tightly coupled with precipitation compared to rainforest NDVI. Rainforest cloud frequency was negatively correlated with the degree of asynchronicity (Delta(NDVI)) between forest types, most strongly at a 1-month lag. Rainforest green-up and dry forest brown-down was particularly apparent during the 2002-003 El Niño. The spatial pattern of NDVI response to the NINO 3.4 Sea Surface Temperature (SST) index during 2002-2003 showed that the leeward side exhibited significant negative correlations to increased SSTs, whereas the windward side exhibited significant positive correlations to increased SSTs, most evident at an 8 to 9-month lag. This study demonstrates that different tropical forest types exhibit asynchronous responses to seasonal and El Niño-driven drought, and suggests that mechanisms controlling dry forest leaf phenology are related to water-limitation, whereas rainforests are more light-limited.

Pan-tropically, liana density increases with decreasing rainfall and increasing seasonality. This pattern has led to the hypothesis that lianas display a growth advantage over trees under dry conditions. However, the physiological mechanisms underpinning this hypothesis remain elusive. A key trait influencing leaf and plant drought tolerance is the leaf water potential at turgor loss point (πtlp). πtlp adjusts under drier conditions and this contributes to improved leaf drought tolerance. For co-occurring Amazonian tree (n = 247) and liana (n = 57) individuals measured during the dry and the wet seasons, lianas showed a stronger osmotic adjustment than trees. Liana leaves were less drought-tolerant than trees in the wet season, but reached similar drought tolerances during the dry season. Stronger osmotic adjustment in lianas would contribute to turgor maintenance, a critical prerequisite for carbon uptake and growth, and to the success of lianas relative to trees in growth under drier conditions.

Background

The extraordinary diversification of angiosperm plants in the Cretaceous and Tertiary periods has produced an estimated 250,000-300,000 living angiosperm species and has fundamentally altered terrestrial ecosystems. Interactions with animals as pollinators or seed dispersers have long been suspected as drivers of angiosperm diversification, yet empirical examples remain sparse or inconclusive. Seed dispersal by ants (myrmecochory) may drive diversification as it can reduce extinction by providing selective advantages to plants and can increase speciation by enhancing geographical isolation by extremely limited dispersal distances.

Methodology/principal findings

Using the most comprehensive sister-group comparison to date, we tested the hypothesis that myrmecochory leads to higher diversification rates in angiosperm plants. As predicted, diversification rates were substantially higher in ant-dispersed plants than in their non-myrmecochorous relatives. Data from 101 angiosperm lineages in 241 genera from all continents except Antarctica revealed that ant-dispersed lineages contained on average more than twice as many species as did their non-myrmecochorous sister groups. Contrasts in species diversity between sister groups demonstrated that diversification rates did not depend on seed dispersal mode in the sister group and were higher in myrmecochorous lineages in most biogeographic regions.

Conclusions/significance

Myrmecochory, which has evolved independently at least 100 times in angiosperms and is estimated to be present in at least 77 families and 11 000 species, is a key evolutionary innovation and a globally important driver of plant diversity. Myrmecochory provides the best example to date for a consistent effect of any mutualism on large-scale diversification.

Ecological niche models (ENMs) provide a means of characterizing the spatial distribution of suitable conditions for species, and have recently been applied to the challenge of locating potential distributional areas at the Last Glacial Maximum (LGM) when unfavorable climate conditions led to range contractions and fragmentation. Here, we compare and contrast ENM-based reconstructions of LGM refugial locations with those resulting from the more traditional molecular genetic and phylogeographic predictions. We examined 20 North American terrestrial vertebrate species from different regions and with different range sizes for which refugia have been identified based on phylogeographic analyses, using ENM tools to make parallel predictions. We then assessed the correspondence between the two approaches based on spatial overlap and areal extent of the predicted refugia. In 14 of the 20 species, the predictions from ENM and predictions based on phylogeographic studies were significantly spatially correlated, suggesting that the two approaches to development of refugial maps are converging on a similar result. Our results confirm that ENM scenario exploration can provide a useful complement to molecular studies, offering a less subjective, spatially explicit hypothesis of past geographic patterns of distribution.

We observed strong positive relationships between soil properties and forest dynamics of growth and mortality across twelve primary lowland tropical forests in a phosphorus-poor region of the Guiana Shield. Average tree growth (diameter at breast height) increased from 0.81 to 2.1 mm yr^-1 along a soil texture gradient from 0 to 67% clay, and increasing metal-oxide content. Soil organic carbon stocks in the top 30 cm ranged from 30 to 118 tons C ha^-1, phosphorus content ranged from 7 to 600 mg kg^-1 soil, and the relative abundance of arbuscular mycorrhizal fungi ranged from 0 to 50%, all positively correlating with soil clay, and iron and aluminum oxide and hydroxide content. In contrast, already low extractable phosphorus (Bray P) content decreased from 4.4 to <0.02 mg kg^-1 in soil with increasing clay content. A greater prevalence of arbuscular mycorrhizal fungi in more clayey forests that had higher tree growth and mortality, but not biomass, indicates that despite the greater investment in nutrient uptake required, soils with higher clay content may actually serve to sustain high tree growth in tropical forests by avoiding phosphorus losses from the ecosystem. Our study demonstrates how variation in soil properties that retain carbon and nutrients can help to explain variation in tropical forest growth and mortality, but not biomass, by requiring niche specialization and contributing to biogeochemical diversification across this region.

Resorption is the active withdrawal of nutrients before leaf abscission. This mechanism represents an important strategy to maintain efficient nutrient cycling; however, resorption is poorly characterized in old-growth tropical forests growing in nutrient-poor soils. We investigated nutrient resorption from leaves in 39 tree species in two tropical forests on the Guiana Shield, French Guiana, to investigate whether resorption efficiencies varied with soil nutrient, seasonality, and species traits. The stocks of P in leaves, litter, and soil were low at both sites, indicating potential P limitation of the forests. Accordingly, mean resorption efficiencies were higher for P (35.9%) and potassium (K; 44.6%) than for nitrogen (N; 10.3%). K resorption was higher in the wet (70.2%) than in the dry (41.7%) season. P resorption increased slightly with decreasing total soil P; and N and P resorptions were positively related to their foliar concentrations. We conclude that nutrient resorption is a key plant nutrition strategy in these old-growth tropical forests, that trees with high foliar nutrient concentration reabsorb more nutrient, and that nutrients resorption in leaves, except P, are quite decoupled from nutrients in the soil. Seasonality and biochemical limitation played a role in the resorption of nutrients in leaves, but species-specific requirements obscured general tendencies at stand and ecosystem level.

The main objective of this paper is to present and discuss the best methods to estimate live above ground biomass in the Atlantic Forest. The methods presented and conclusions are the products of a workshop entitled "Estimation of Biomass and Carbon Stocks: the Case of Atlantic Rain Forest". Aboveground biomass (AGB) in tropical forests is mainly contained in trees. Tree biomass is a function of wood volume, obtained from the diameter and height, architecture and wood density (dry weight per unit volume of fresh wood). It can be quantified by the direct (destructive) or indirect method where the biomass quantification is estimated using mathematical models. The allometric model can be site specific when elaborated to a particular ecosystem or general that can be used in different sites. For the Atlantic Forest, despite the importance of it, there are only two direct measurements of tree biomass, resulting in allometric models specific for this ecosystem. To select one or other of the available models in the literature to estimate AGB it is necessary take into account what is the main question to be answered and the ease with which it is possible to measure the independent variables in the model. Models that present more accurate estimates should be preferred. However, more simple models (those with one independent variable, usually DBH) can be used when the focus is monitoring the variation in carbon storage through the time. Our observations in the Atlantic Forest suggest that pan-tropical relations proposed by Chave et al. (2005) can be confidently used to estimated tree biomass across biomes as long as tree diameter (DBH), height, and wood density are accounted for in the model. In Atlantic Forest, we recommend the quantification of biomass of lianas, bamboo, palms, tree ferns and epiphytes, which are an important component in this ecosystem. This paper is an outcome of the workshop entitled "Estimation of Biomass and Carbon Stocks: the Case of Atlantic Rain Forest", that was conducted at Ubatuba, São Paulo, Brazil, between 4 and 8 December 2006 as part of the Brazilian project "Ombrophylus Dense Forest floristic composition, structure and function at the Núcleos Picinguaba and Santa Virginia of the Serra do Mar State Park", BIOTA Gradiente.

The net primary production of tropical forests and its partitioning between long-lived carbon pools (wood) and shorter-lived pools (leaves, fine roots) are of considerable importance in the global carbon cycle. However, these terms have only been studied at a handful of field sites, and with no consistent calculation methodology. Here we calculate above-ground coarse wood carbon productivity for 104 forest plots in lowland New World humid tropical forests, using a consistent calculation methodology that incorporates corrections for spatial variations in tree-size distributions and wood density, and for census interval length. Mean wood density is found to be lower in more productive forests. We estimate that above-ground coarse wood productivity varies by more than a factor of three (between 1.5 and 5.5 Mg C ha-1a-1) across the Neotropical plots, with a mean value of 3.1 Mg C ha-a-1. There appear to be no obvious relationships between wood productivity and rainfall, dry season length or sunshine, but there is some hint of increased productivity at lower temperatures. There is, however, also strong evidence for a positive relationship between wood productivity and soil fertility. Fertile soils tend to become more common towards the Andes and at slightly higher than average elevations, so the apparent temperature/productivity relationship is probably not a direct one. Coarse wood productivity accounts for only a fraction of overall tropical forest net primary productivity, but the available data indicate that it is approximately proportional to total above-ground productivity. We speculate that the large variation in wood productivity is unlikely to directly imply an equivalent variation in gross primary production. Instead a shifting balance in carbon allocation between respiration, wood carbon and fine root production seems the more likely explanation. © 2004 Blackwell Publishing Ltd.

Remote sensing is revolutionizing the way we study forests, and recent technological advances mean we are now able - for the first time - to identify and measure the crown dimensions of individual trees from airborne imagery. Yet to make full use of these data for quantifying forest carbon stocks and dynamics, a new generation of allometric tools which have tree height and crown size at their centre are needed. Here, we compile a global database of 108753 trees for which stem diameter, height and crown diameter have all been measured, including 2395 trees harvested to measure aboveground biomass. Using this database, we develop general allometric models for estimating both the diameter and aboveground biomass of trees from attributes which can be remotely sensed - specifically height and crown diameter. We show that tree height and crown diameter jointly quantify the aboveground biomass of individual trees and find that a single equation predicts stem diameter from these two variables across the world's forests. These new allometric models provide an intuitive way of integrating remote sensing imagery into large-scale forest monitoring programmes and will be of key importance for parameterizing the next generation of dynamic vegetation models.