Coexistence and environmental filtering of species-specific biomass in an African savanna.

Biomass density is a key metric of vegetation abundance, but understanding how community assembly processes, such as environmental filtering and competitive exclusion, affect biomass distributions of coexisting species has proven logistically challenging. Here we apply airborne remote sensing to study the ecosystem-scale distribution of species-specific, woody plant biomass and its relation to topographic and hydrologic gradients in a South African savanna. We also spatially analyzed variation in biomass among species to understand patterns of coexistence, mapping the species and biomass over one million trees across 10500 ha. We found the biomass of dominant woody species to be weakly but significantly related to environmental filters, where a combination of 10 topographic and edaphic variables accounted for < 15% of the variance in the biomass of any given species. Distance to nearest stream was the only environmental variable significantly correlated to all species' biomass. Despite an overall negative trend observed between the biomass of species pairs, we found a number of regions where the biomass of two species was similar or equal, and all species pairs exhibited some level of co-occurrence. This suggests that even weak stabilizing mechanisms (e.g., environmental niches) can overcome fitness differences and balance competitive exclusion, enabling coexistence. Future work of repeated measurements of species-specific biomass will provide a novel advance in understanding woody plant community assembly processes in natural ecosystems. Characterizing the species composition of biomass is an important advance in understanding the balance of community assembly processes and its control over current species assemblages.

Improved allometric models to estimate the aboveground biomass of tropical trees.

Terrestrial carbon stock mapping is important for the successful implementation of climate change mitigation policies. Its accuracy depends on the availability of reliable allometric models to infer oven-dry aboveground biomass of trees from census data. The degree of uncertainty associated with previously published pantropical aboveground biomass allometries is large. We analyzed a global database of directly harvested trees at 58 sites, spanning a wide range of climatic conditions and vegetation types (4004 trees ≥ 5 cm trunk diameter). When trunk diameter, total tree height, and wood specific gravity were included in the aboveground biomass model as covariates, a single model was found to hold across tropical vegetation types, with no detectable effect of region or environmental factors. The mean percent bias and variance of this model was only slightly higher than that of locally fitted models. Wood specific gravity was an important predictor of aboveground biomass, especially when including a much broader range of vegetation types than previous studies. The generic tree diameter-height relationship depended linearly on a bioclimatic stress variable E, which compounds indices of temperature variability, precipitation variability, and drought intensity. For cases in which total tree height is unavailable for aboveground biomass estimation, a pantropical model incorporating wood density, trunk diameter, and the variable E outperformed previously published models without height. However, to minimize bias, the development of locally derived diameter-height relationships is advised whenever possible. Both new allometric models should contribute to improve the accuracy of biomass assessment protocols in tropical vegetation types, and to advancing our understanding of architectural and evolutionary constraints on woody plant development.

Harvesting tree biomass at the stand level to assess the accuracy of field and airborne biomass estimation in savannas.

Tree biomass is an integrated measure of net growth and is critical for understanding, monitoring, and modeling ecosystem functions. Despite the importance of accurately measuring tree biomass, several fundamental barriers preclude direct measurement at large spatial scales, including the facts that trees must be felled to be weighed and that even modestly sized trees are challenging to maneuver once felled. Allometric methods allow for estimation of tree mass using structural characteristics, such as trunk diameter. Savanna trees present additional challenges, including limited available allometry and a prevalence of multiple stems per individual. Here we collected airborne lidar data over a semiarid savanna adjacent to the Kruger National Park, South Africa, and then harvested and weighed woody plant biomass at the plot scale to provide a standard against which field and airborne estimation methods could be compared. For an existing airborne lidar method, we found that half of the total error was due to averaging canopy height at the plot scale. This error was eliminated by instead measuring maximum height and crown area of individual trees from lidar data using an object-based method to identify individual tree crowns and estimate their biomass. The best object-based model approached the accuracy of field allometry at both the tree and plot levels, and it more than doubled the accuracy compared to existing airborne methods (17% vs. 44% deviation from harvested biomass). Allometric error accounted for less than one-third of the total residual error in airborne biomass estimates at the plot scale when using allometry with low bias. Airborne methods also gave more accurate predictions at the plot level than did field methods based on diameter-only allometry. These results provide a novel comparison of field and airborne biomass estimates using harvested plots and advance the role of lidar remote sensing in savanna ecosystems.

Large-scale impacts of herbivores on the structural diversity of African savannas.

African savannas are undergoing management intensification, and decision makers are increasingly challenged to balance the needs of large herbivore populations with the maintenance of vegetation and ecosystem diversity. Ensuring the sustainability of Africa's natural protected areas requires information on the efficacy of management decisions at large spatial scales, but often neither experimental treatments nor large-scale responses are available for analysis. Using a new airborne remote sensing system, we mapped the three-dimensional (3-D) structure of vegetation at a spatial resolution of 56 cm throughout 1640 ha of savanna after 6-, 22-, 35-, and 41-year exclusions of herbivores, as well as in unprotected areas, across Kruger National Park in South Africa. Areas in which herbivores were excluded over the short term (6 years) contained 38%-80% less bare ground compared with those that were exposed to mammalian herbivory. In the longer-term (> 22 years), the 3-D structure of woody vegetation differed significantly between protected and accessible landscapes, with up to 11-fold greater woody canopy cover in the areas without herbivores. Our maps revealed 2 scales of ecosystem response to herbivore consumption, one broadly mediated by geologic substrate and the other mediated by hillslope-scale variation in soil nutrient availability and moisture conditions. Our results are the first to quantitatively illustrate the extent to which herbivores can affect the 3-D structural diversity of vegetation across large savanna landscapes.

Tree foliar chemistry in an African savanna and its relation to life history strategies and environmental filters.

Understanding the relative importance of environment and life history strategies in determining leaf chemical traits remains a key objective of plant ecology. We assessed 20 foliar chemical properties among 12 African savanna woody plant species and their relation to environmental variables (hillslope position, precipitation, geology) and two functional traits (thorn type and seed dispersal mechanism). We found that combinations of six leaf chemical traits (lignin, hemi-cellulose, zinc, boron, magnesium, and manganese) predicted the species with 91% accuracy. Hillslope position, precipitation, and geology accounted for only 12% of the total variance in these six chemical traits. However, thorn type and seed dispersal mechanism accounted for 46% of variance in these chemical traits. The physically defended species had the highest concentrations of hemi-cellulose and boron. Species without physical defense had the highest lignin content if dispersed by vertebrates, but threefold lower lignin content if dispersed by wind. One of the most abundant woody species in southern Africa, Colophospermum mopane, was found to have the highest foliar concentrations of zinc, phosphorus, and δ(13)C, suggesting that zinc chelation may be used by this species to bind metallic toxins and increase uptake of soil phosphorus. Across all studied species, taxonomy and physical traits accounted for the majority of variability in leaf chemistry.

Human and environmental controls over aboveground carbon storage in Madagascar.

BACKGROUND: Accurate, high-resolution mapping of aboveground carbon density (ACD, Mg C ha-1) could provide insight into human and environmental controls over ecosystem state and functioning, and could support conservation and climate policy development. However, mapping ACD has proven challenging, particularly in spatially complex regions harboring a mosaic of land use activities, or in remote montane areas that are difficult to access and poorly understood ecologically. Using a combination of field measurements, airborne Light Detection and Ranging (LiDAR) and satellite data, we present the first large-scale, high-resolution estimates of aboveground carbon stocks in Madagascar. RESULTS: We found that elevation and the fraction of photosynthetic vegetation (PV) cover, analyzed throughout forests of widely varying structure and condition, account for 27-67% of the spatial variation in ACD. This finding facilitated spatial extrapolation of LiDAR-based carbon estimates to a total of 2,372,680 ha using satellite data. Remote, humid sub-montane forests harbored the highest carbon densities, while ACD was suppressed in dry spiny forests and in montane humid ecosystems, as well as in most lowland areas with heightened human activity. Independent of human activity, aboveground carbon stocks were subject to strong physiographic controls expressed through variation in tropical forest canopy structure measured using airborne LiDAR. CONCLUSIONS: High-resolution mapping of carbon stocks is possible in remote regions, with or without human activity, and thus carbon monitoring can be brought to highly endangered Malagasy forests as a climate-change mitigation and biological conservation strategy.