ALBERT

Description: Leaves in tropical forests come in an enormous variety of sizes and shapes, each of which can be ultimately viewed as an adaptation to the complex problem of optimising the capture of light for photosynthesis. However, the fact that many different shape "strategies" coexist within a habitat demonstrate that there are many other intrinsic and extrinsic factors involved, such as the differential investment in support tissues required for different leaf lamina shapes. Here, we take a macrogeographic approach to understanding the function of different lamina shape categories. Specifically, we use 106 permanent plots spread across the Amazon rainforest basin to: 1) describe the geographic distribution of some simple metrics of lamina shape in plots from across Amazonia, and; 2) identify and quantify relationships between key environmental parameters and lamina shape in tropical forests. Because the plots are not randomly distributed across the study area, achieving this latter objective requires the use of statistics that can account for spatial auto-correlation. We found that between 60–70% of the 2791 species and 83 908 individual trees in the dataset could be classified as having elliptic leaves (= the widest part of the leaf is on an axis in the middle fifth of the long axis of the leaf). Furthermore, the average Amazonian tree leaf is 2.5 times longer than it is wide and has an entire margin. Contrary to theoretical expectations we found little support for the hypothesis that narrow leaves are an adaptation to dry conditions. However, we did find strong regional patterns in leaf lamina length-width ratios and several significant correlations with precipitation variables suggesting that water availability may be exerting an as yet unrecognised selective pressure on leaf shape of rainforest trees. Some support was found for the hypothesis that narrow leaves are an adaptation to low nutrient soils. Furthermore, we found a strong correlation between the proportion of trees with non-entire laminas (dissected, toothed, etc.) and mean annual temperature once again supporting the well documented association that provides a basis for reconstructing past temperature regimes.

Description: Linking lower and higher trophic levels requires special focus on the essential role played by mid-trophic levels, i.e., the zooplankton. One of the most relevant pieces of information regarding zooplankton in terms of flux of energy lies in its size structure. In this study, an extensive data set of size measurements is presented, covering parts of the western European continental shelf and slope, from the Galician coast to the Ushant front, during the springs from 2005 to 2012. Zooplankton size spectra were estimated using measurements carried out in situ with the Laser Optical Plankton Counter (LOPC) and with an image analysis of WP2 net samples (200 μm mesh size) performed following the ZooScan methodology. The LOPC counts and sizes particles within 100–2000 μm of spherical equivalent diameter (ESD), whereas the WP2/ZooScan allows for counting, sizing and identification of zooplankton from ~ 400 μm ESD. The difference between the LOPC (all particles) and the WP2/ZooScan (zooplankton only) was assumed to provide the size distribution of non-living particles, whose descriptors were related to a set of explanatory variables (including physical, biological and geographic descriptors). A statistical correction based on these explanatory variables was further applied to the LOPC size distribution in order to remove the non-living particles part, and therefore estimate the size distribution of zooplankton. This extensive data set provides relevant information about the zooplankton size distribution variability, productivity and trophic transfer efficiency in the pelagic ecosystem of the Bay of Biscay at a regional and interannual scale.

Description: Linking lower to higher trophic levels requires a special focus on the pivotal role played by mid-trophic levels, i.e. the zooplankton. One of the most relevant information on zooplankton in term of fluxes of matter lies in its size structure. We present here an extensive dataset of size measurements covering part of the western European shelf and slope, from the Galician coast to the Ushant front, during springs from 2005 to 2012. Zooplankton size spectra were estimated using both measurements carried out in situ by the Laser-Optical Plankton Counter (LOPC, 816 records) and WP2 net (200 μm mesh size) samples scanned following the ZooScan methodology and image analysis (a total of 89 samples were analyzed). The LOPC counts and sizes all particles in the range 100 to 2000 μm of spherical equivalent diameter (ESD) whereas the WP2/ZooScan allows the counting, sizing and identification of zooplankton from ~400 μm ESD. The difference between the LOPC (all particles) and the WP2/ZooScan (zooplankton only) is assumed to provide the size distribution of non-living particles whose descriptors are further related to a set of explanatory variables (including physical, biological and geographic descriptors). A statistical correction based on these explanatory variables is then applied to LOPC measurements to removed the part due to non-living particles and estimate zooplankton size spectra. This extensive data set provides a new look at regional and inter-annual variability of the pelagic ecosystem of the Bay of Biscay.

Description: Leaves in tropical forests come in an enormous variety of sizes and shapes, each of which can be ultimately viewed as an adaptation to the complex problem of optimising the capture of light for photosynthesis. However, the fact that many different shape "strategies" coexist within a habitat demonstrate that there are many other intrinsic and extrinsic factors involved, such as the differential investment in support tissues required for different leaf lamina shapes. Here, we take a macrogeographic approach to understanding the function of different lamina shape categories. Specifically, we use 106 permanent plots spread across the Amazon rainforest basin to: (1) describe the geographic distribution of some simple metrics of lamina shape in plots from across Amazonia, and; (2) identify and quantify relationships between key environmental parameters and lamina shape in tropical forests. Because the plots are not randomly distributed across the study area, achieving this latter objective requires the use of statistics that can account for spatial auto-correlation. We found that between 60–70% of the 2791 species and 83 908 individual trees in the dataset could be classified as elliptic (=the widest part of a leaf is on an axis in the middle fifth of the long axis of the leaf). Furthermore, the average Amazonian tree leaf is 2.5 times longer than it is wide and has an entire margin. Contrary to theoretical expectations we found little support for the hypothesis that narrow leaves are an adaptation to dry conditions and low nutrient soils. However, we did find strong regional patterns in leaf lamina length-width ratios and several significant correlations with precipitation variables suggesting that water availability may be exerting an as yet unrecognised selective pressure on leaf shape of rainforest trees. Furthermore, we found a strong correlation between the proportion of trees with non-entire laminas (dissected, toothed, etc.) and mean annual temperature once again supporting the well documented association that provides a basis for reconstructing past temperature regimes.

Description: Leaf size influences many aspects of tree function such as rates of transpiration and photosynthesis and, consequently, often varies in a predictable way in response to environmental gradients. The recent development of pan-Amazonian databases based on permanent botanical plots (e.g. RAINFOR, ATDN) has now made it possible to assess trends in leaf size across environmental gradients in Amazonia. Previous plot-based studies have shown that the community structure of Amazonian trees breaks down into at least two major ecological gradients corresponding with variations in soil fertility (decreasing south to northeast) and length of the dry season (increasing from northwest to south and east). Here we describe the results of the geographic distribution of leaf size categories based on 121 plots distributed across eight South American countries. We find that, as predicted, the Amazon forest is predominantly populated by tree species and individuals in the mesophyll size class (20.25–182.25 cm2). The geographic distribution of species and individuals with large leaves (〉20.25 cm2) is complex but is generally characterized by a higher proportion of such trees in the north-west of the region. Spatially corrected regressions reveal weak correlations between the proportion of large-leaved species and metrics of water availability. We also find a significant negative relationship between leaf size and wood density.

Description: Tropical tree height-diameter (H:D) relationships may vary by forest type and region making large-scale estimates of above-ground biomass subject to bias if they ignore these differences in stem allometry. We have therefore developed a new global tropical forest database consisting of 39 955 concurrent H and D measurements encompassing 283 sites in 22 tropical countries. Utilising this database, our objectives were:   1. to determine if H:D relationships differ by geographic region and forest type (wet to dry forests, including zones of tension where forest and savanna overlap).   2. to ascertain if the H:D relationship is modulated by climate and/or forest structural characteristics (e.g. stand-level basal area, A).   3. to develop H:D allometric equations and evaluate biases to reduce error in future local-to-global estimates of tropical forest biomass. Annual precipitation coefficient of variation (PV), dry season length (SD), and mean annual air temperature (TA) emerged as key drivers of variation in H:D relationships at the pantropical and region scales. Vegetation structure also played a role with trees in forests of a high A being, on average, taller at any given D. After the effects of environment and forest structure are taken into account, two main regional groups can be identified. Forests in Asia, Africa and the Guyana Shield all have, on average, similar H:D relationships, but with trees in the forests of much of the Amazon Basin and tropical Australia typically being shorter at any given D than their counterparts elsewhere. The region-environment-structure model with the lowest Akaike's information criterion and lowest deviation estimated stand-level H across all plots to within a median –2.7 to 0.9% of the true value. Some of the plot-to-plot variability in H:D relationships not accounted for by this model could be attributed to variations in soil physical conditions. Other things being equal, trees tend to be more slender in the absence of soil physical constraints, especially at smaller D. Pantropical and continental-level models provided only poor estimates of H, especially when the roles of climate and stand structure in modulating H:D allometry were not simultaneously taken into account.

Description: Above-ground tropical tree biomass and carbon storage estimates commonly ignore tree height. We estimate the effect of incorporating height (H) on forest biomass estimates using 37 625 concomitant H and diameter measurements (n = 327 plots) and 1816 harvested trees (n = 21 plots) tropics-wide to answer the following questions: 1. For trees of known biomass (from destructive harvests) which H-model form and geographic scale (plot, region, and continent) most reduces biomass estimate uncertainty? 2. How much does including H relationship estimates derived in (1) reduce uncertainty in biomass estimates across 327 plots spanning four continents? 3. What effect does the inclusion of H in biomass estimates have on plot- and continental-scale forest biomass estimates? The mean relative error in biomass estimates of the destructively harvested trees was half (mean 0.06) when including H, compared to excluding H (mean 0.13). The power- and Weibull-H asymptotic model provided the greatest reduction in uncertainty, with the regional Weibull-H model preferred because it reduces uncertainty in smaller-diameter classes that contain the bulk of biomass per hectare in most forests. Propagating the relationships from destructively harvested tree biomass to each of the 327 plots from across the tropics shows errors are reduced from 41.8 Mg ha−1 (range 6.6 to 112.4) to 8.0 Mg ha−1 (−2.5 to 23.0) when including $H$. For all plots, above-ground live biomass was 52.2±17.3 Mg ha−1 lower when including H estimates (13%), with the greatest reductions in estimated biomass in Brazilian Shield forests and relatively no change in the Guyana Shield, central Africa and southeast Asia. We show fundamentally different stand structure across the four forested tropical continents, which affects biomass reductions due to $H$. African forests store a greater portion of total biomass in large-diameter trees and trees are on average larger in diameter. This contrasts to forests on all other continents where smaller-diameter trees contain the greatest fractions of total biomass. After accounting for variation in $H$, total biomass per hectare is greatest in Australia, the Guyana Shield, and Asia and lowest in W. Africa, W. Amazonia, and the Brazilian Shield (descending order). Thus, if closed canopy tropical forests span 1668 million km2 and store 285 Pg C, then the overestimate is 35 Pg C if H is ignored, and the sampled plots are an unbiased statistical representation of all tropical forest in terms of biomass and height factors. Our results show that tree $H$ is an important allometric factor that needs to be included in future forest biomass estimates to reduce error in estimates of pantropical carbon stocks and emissions due to deforestation.

Description: Landscape and environmental variables such as topography, geomorphology, soil types, and climate are important factors affecting forest composition, structure, productivity, and biomass. Here, we combine a network of forest inventories with recently developed global data products from satellite observations in modeling the potential distributions of forest structure and productivity in Amazonia and examine how geomorphology, soil, and precipitation control these distributions. We use the RAINFOR network of forest plots distributed in lowland forests across Amazonia, and satellite observations of tree cover, leaf area index, phenology, moisture, and topographical variations. A maximum entropy estimation (Maxent) model is employed to predict the spatial distribution of several key forest structure parameters: basal area, fraction of large trees, fraction of palms, wood density, productivity, and above-ground biomass at 5 km spatial resolution. A series of statistical tests at selected thresholds as well as across all thresholds and jackknife analysis are used to examine the accuracy of distribution maps and the relative contributions of environmental variables. The final maps were interpreted using soil, precipitation, and geomorphological features of Amazonia and it was found that the length of dry season played a key role in impacting the distribution of all forest variables except the wood density. Soil type had a significant impact on the wood productivity. Most high productivity forests were distributed either on less infertile soils of western Amazonia and Andean foothills, on crystalline shields, and younger alluvial deposits. Areas of low elevation and high density of small rivers of Central Amazonia showed distinct features, hosting mainly forests with low productivity and smaller trees.

Description: Long-term studies using the RAINFOR network of forest plots have generated significant insights into the spatial and temporal dynamics of forest carbon cycling in Amazonia. In this work, we map and explore the landscape context of several major RAINFOR plot clusters using Landsat ETM+ satellite data. In particular, we explore how representative the plots are of their landscape context, and test whether bias in plot location within landscapes may be influencing the regional mean values obtained for important forest biophysical parameters. Specifically, we evaluate whether the regional variations in wood productivity, wood specific density and above ground biomass derived from the RAINFOR network could be driven by systematic and unintentional biases in plot location. Remote sensing data covering 45 field plots were aggregated to generate landscape maps to identify the specific physiognomy of the plots. In the Landsat ETM+ data, it was possible to spectrally differentiate three types of terra firme forest, three types of alluvial terrain forest, two types of bamboo-dominated forest, palm forest, Heliconia monodominant vegetation, swamp forest, disturbed forests and land use areas. Overall, the plots were generally representative of the forest physiognomies in the landscape in which they are located. Furthermore, the analysis supports the observed regional trends in those important forest parameters. This study demonstrates the utility of landscape scale analysis of forest physiognomies for validating and supporting the finds of plot based studies. Moreover, the more precise geolocation of many key RAINFOR plot clusters achieved during this research provides important contextual information for studies employing the RAINFOR database.

Description: The Amazon basin, one of the most substantial biomass carbon pools on earth, is characterised by strong macroecological gradients in biomass, mortality rates, and wood density from the west to the east. These gradients could affect necromass stocks, but this has not yet been tested. This study aims to assess the stocks and determinants of necromass patterns across Amazonian forests. Field-based and literature data were used to find relationships between necromass and possible determinants. The final regression result was used to estimate and extrapolate the necromass stocks across terra firma Amazonian forests. In eight northwestern and three northeastern Amazonian permanent plots, volumes of coarse woody debris (≥10 cm diameter) were measured in the field and density of each decay class was estimated. Forest structure and historical mortality data were used to determine controlling factors of necromass. Necromass is greater in forests with low stem mortality rates (northeast) rather than forest with high stem mortality rates (northwest) (58.5±10.6 and 27.3±3.2 Mg ha−1, respectively). After integrating all published necromass values, we find that necromass across terra firma forests in Amazonia is positively related to stand biomass, mortality mass input, and average wood density of live trees (ρBA j). We applied these relationships to estimate necromass for plots where necromass has not been measured. The estimates, together with other actual measurements of necromass, were scaled-up to project a total Amazonian necromass of 9.6±1.0 Pg C. The ratio of necromass (on average weighted by forest region) to coarse aboveground biomass is 0.127. Overall, we find (1) a strong spatial trend in necromass in parallel with other macroecological gradients and (2) that necromass is a substantial component of the carbon pool in the Amazon.