Tropical Savanna Ecosystems Research Articles

Does long‐term fire exclusion in an Australian tropical savanna result in a biome shift? A test using the reintroduction of fire

AbstractThe structure of tropical savanna ecosystems is influenced by fire frequency and intensity. There is particular interest in the extent to which long‐term fire exclusion can result in a shift from savanna to forest vegetation that is not easily reversed by the reintroduction of fire. This study examined changes in the structure and composition of a long‐unburnt site within the northern Australian savannas following an extended period of active fire exclusion (>20 years), and the effect of the reintroducing fire through experimental fire regimes, including fires in the early and late dry season at a range of frequencies. After the long period of fire exclusion, the vegetation community was characterized by a well‐developed midstorey and canopy layer, low grass cover, substantially higher densities of woody sprouts and saplings than frequently burnt savanna. The community composition included a high proportion of rainforest‐affiliated species. Three years of experimental fires had no detectable effect on the overall composition of grass layer and woody plants but had an effect on woody vegetation structure. Continued fire exclusion further increased the density of woody stems, particularly in the midstorey (2.0–4.99 m), whereas moderate‐intensity fires (>800 kW m−1) significantly reduced the density of midstorey stems. The reintroduction of higher moderate intensity fire events resulted in the vegetation in some compartments reverting to the open savanna structure typical of frequently burnt sites. Such rapid reversibility suggests that in general, the woody thickening resulting from long‐term fire exclusion did not represent a biome shift to a non‐savanna state. However, there was a small proportion of the site that could not sustain the fires applied to them because grass cover was very low and patchy and therefore appeared to have crossed an ecological threshold towards closed forest.

Soil particle accumulation in termite (Macrotermes bellicosus) mounds and the implications for soil particle dynamics in a tropical savanna Ultisol

AbstractThis study investigated the influence of mound‐building termites on soil particle dynamics on the land surface and in soil‐forming processes by examining the amount of soil particles in mound structures of Macrotermes bellicosus in a highly weathered Ultisol of tropical savanna. Soil particle turnover via the mounds was estimated using particle stock data and soil turnover data from previous studies. A 4‐ha study plot with six mounds of relatively uniform shape and size was investigated. Soil mass constituting the mounds was 6,166 ± 1,581 kg mound−1 within which the mound wall and nest body accounted for 5,002 ± 1,289 and 1,164 ± 293 kg, respectively. The mound wall contained a significantly larger amount of clay (252 ± 9.97 g kg−1) balanced with a lower sand content (676 ± 26.5 g kg−1) than in the adjacent surface (Ap1) horizon, (46.4 ± 12.8 g clay kg−1; 866 ± 83.2 g sand kg−1); the nest body had much higher clay content (559 ± 51.0 g kg−1) but less sand (285 ± 79.2 g kg−1) than the mound wall. As a result, the mounds of M. bellicosus accumulated clay of 2,874 ± 781 kg ha−1 (corresponding to 2.52% of clay stock in the Ap1 horizon) along with an estimated clay turnover rate of 169 kg ha−1 year−1. These findings suggest a positive feedback effect from termite mound‐building activity on soil particle dynamics in tropical savanna ecosystems: M. bellicosus preferentially use subsoil material for mound construction, resulting in relocation of illuvial clay in the subsoil to the land surface where clay eluviation from the surface soil and its illuviation in the subsoil are major soil‐forming processes.

Seasonal variation and fire effects on CH4, N2O and CO2 exchange in savanna soils of northern Australia

Tropical savanna ecosystems are a major contributor to global CO2, CH4 and N2O greenhouse gas exchange. Savanna fire events represent large, discrete C emissions but the importance of ongoing soil-atmosphere gas exchange is less well understood. Seasonal rainfall and fire events are likely to impact upon savanna soil microbial processes involved in N2O and CH4 exchange. We measured soil CO2, CH4 and N2O fluxes in savanna woodland (Eucalyptus tetrodonta/Eucalyptus miniata trees above sorghum grass) at Howard Springs, Australia over a 16 month period from October 2007 to January 2009 using manual chambers and a field-based gas chromatograph connected to automated chambers. The effect of fire on soil gas exchange was investigated through two controlled burns and protected unburnt areas. Fire is a frequent natural and management action in these savanna (every 1–2 years). There was no seasonal change and no fire effect upon soil N2O exchange. Soil N2O fluxes were very low, generally between −1.0 and 1.0μg Nm−2h−1, and often below the minimum detection limit. There was an increase in soil NH4+ in the months after the 2008 fire event, but no change in soil NO3−. There was considerable nitrification in the early wet season but minimal nitrification at all other times.Savanna soil was generally a net CH4 sink that equated to between −2.0 and −1.6kg CH4ha−1y−1 with no clear seasonal pattern in response to changing soil moisture conditions. Irrigation in the dry season significantly reduced soil gas diffusion and as a consequence soil CH4 uptake. There were short periods of soil CH4 emission, up to 20μg Cm−2h−1, likely to have been caused by termite activity in, or beneath, automated chambers. Soil CO2 fluxes showed a strong bimodal seasonal pattern, increasing fivefold from the dry into the wet season. Soil moisture showed a weak relationship with soil CH4 fluxes, but a much stronger relationship with soil CO2 fluxes, explaining up to 70% of the variation in unburnt treatments. Australian savanna soils are a small N2O source, and possibly even a sink. Annual soil CH4 flux measurements suggest that the 1.9million km2 of Australian savanna soils may provide a C sink of between −7.7 and −9.4 Tg CO2-e per year. This sink estimate would offset potentially 10% of Australian transport related CO2-e emissions. This CH4 sink estimate does not include concurrent CH4 emissions from termite mounds or ephemeral wetlands in Australian savannas.

Moisture can be the dominant environmental parameter governing cadaver decomposition in soil

Forensic taphonomy involves the use of decomposition to estimate postmortem interval (PMI) or locate clandestine graves. Yet, cadaver decomposition remains poorly understood, particularly following burial in soil. Presently, we do not know how most edaphic and environmental parameters, including soil moisture, influence the breakdown of cadavers following burial and alter the processes that are used to estimate PMI and locate clandestine graves. To address this, we buried juvenile rat ( Rattus rattus) cadavers (∼18 g wet weight) in three contrasting soils from tropical savanna ecosystems located in Pallarenda (sand), Wambiana (medium clay), or Yabulu (loamy sand), Queensland, Australia. These soils were sieved (2 mm), weighed (500 g dry weight), calibrated to a matric potential of -0.01 megapascals (MPa), -0.05 MPa, or -0.3 MPa (wettest to driest) and incubated at 22 °C. Measurements of cadaver decomposition included cadaver mass loss, carbon dioxide-carbon (CO 2-C) evolution, microbial biomass carbon (MBC), protease activity, phosphodiesterase activity, ninhydrin-reactive nitrogen (NRN) and soil pH. Cadaver burial resulted in a significant increase in CO 2-C evolution, MBC, enzyme activities, NRN and soil pH. Cadaver decomposition in loamy sand and sandy soil was greater at lower matric potentials (wetter soil). However, optimal matric potential for cadaver decomposition in medium clay was exceeded, which resulted in a slower rate of cadaver decomposition in the wettest soil. Slower cadaver decomposition was also observed at high matric potential (-0.3 MPa). Furthermore, wet sandy soil was associated with greater cadaver decomposition than wet fine-textured soil. We conclude that gravesoil moisture content can modify the relationship between temperature and cadaver decomposition and that soil microorganisms can play a significant role in cadaver breakdown. We also conclude that soil NRN is a more reliable indicator of gravesoil than soil pH.

Mound building termites contribute to savanna vegetation heterogeneity

With biomass densities comparable to large ungulates and megaherbivores, termites play a key functional role in many tropical savanna ecosystems. This study focuses on vegetated termite mounds (termitaria) constructed by the Termitidae species Macrotermes herus. We studied how resource rich termitaria affect graminoid herbs (Poaceae and Cyperaceae), forbs and woody species composition and diversity. The density of termitaria explained 89% of the variation in dense thickets in the area. Fire tolerant Acacia species dominated the open savanna while fire sensitive species like Grewia spp. and the succulent Euphorbia candelabrum were restricted to termite mounds. Termitaria plots had four times the mean number of woody species and supported three times as many forb species as the adjacent savanna. For woody species, both the Shannon–Wiener index and the Shannon evenness index were higher on temitaria than on the savanna. There were no differences for graminoid herbs, except for the Shannon evenness index which was higher on termitaria. Our results indicate that graminoid herb richness peaks at lower productivity levels than trees and forbs in savanna ecosystems, as also recently found in temperate areas.

Tropical forest and savanna ecosystems show differential impact of N and P additions on soil organic matter and aggregate structure

AbstractLandscape transformation and atmospheric nutrient depositions, important global change drivers, are affecting the vegetation and soil properties of natural dry tropical forest and derived savanna ecosystems in India. This study assessed the effect of continuous N and P additions for 6 years on the size distribution and properties of soil aggregates in forest–ecotone–savanna gradient. Addition of N significantly increased the proportion of macroaggregates in forest and ecotone, whereas the same input significantly decreased their proportion in the savanna. Consequently, the stability of soil aggregates increased significantly in forest and ecotone, whereas it decreased significantly in the savanna. The effect of P addition on soil aggregate stability was marginal. N addition also altered the biological and chemical qualities of soil aggregates. It caused increase in microbial biomass C (MBC) associated with macroaggregates in forest and ecotone; however, in savanna, MBC increased in the microaggregates. P addition did not affect the amount of MBC in both types of soil aggregates. Because of rapid accumulation of applied N and P in the microbial biomass, the ratios of MBC to microbial biomass nitrogen (MBN) as well as microbial biomass phosphorous (MBP) were decreased in both aggregates. Overall, the effect of N addition was more marked than that of P addition, suggesting that N is more limiting than P in these dry tropical ecosystems. In the current scenario of N loading, continued soil N loading in forest may lead to increased macroaggregates with associated MBC and MBN and greater aggregate stability. In contrast, the extensively distributed savannas may show the reverse trend leading to a decrease in soil fertility.

Temperature affects microbial decomposition of cadavers ( Rattus rattus) in contrasting soils

The ecology of soils associated with dead mammals (i.e. cadavers) is poorly understood. Although temperature and soil type are well known to influence the decomposition of other organic resource patches, the effect of these variables on the degradation of cadavers in soil has received little experimental investigation. To address this, cadavers of juvenile rats ( Rattus rattus) were buried in one of three contrasting soils (Sodosol, Rudosol, and Vertosol) from tropical savanna ecosystems in Queensland, Australia and incubated at 29 °C, 22 °C, or 15 °C in a laboratory setting. Cadavers and soils were destructively sampled at intervals of 7 days over an incubation period of 28 days. Measurements of decomposition included cadaver mass loss, carbon dioxide–carbon (CO 2–C) evolution, microbial biomass carbon (MBC), protease activity, phosphodiesterase activity, and soil pH, which were all significantly positively affected by cadaver burial. A temperature effect was observed where peaks or differences in decomposition that at occurred at higher temperature would occur at later sample periods at lower temperature. Soil type also had an important effect on some measured parameters. These findings have important implications for a largely unexplored area of soil ecology and nutrient cycling, which are significant for forensic science, cemetery planning and livestock carcass disposal.

Nutrient resorption patterns of plant functional groups in a tropical savanna: variation and functional significance

Green and senesced leaf nitrogen (N) and phosphorus (P) concentrations of different plant functional groups in savanna communities of Kruger National Park, South Africa were analyzed to determine if nutrient resorption was regulated by plant nutritional status and foliar N:P ratios. The N and P concentrations in green leaves and the N concentrations in senesced leaves differed significantly between the dominant plant functional groups in these savannas: fine-leaved trees, broad-leaved trees and grasses. However, all three functional groups reduced P to comparable and very low levels in senesced leaves, suggesting that P was tightly conserved in this tropical semi-arid savanna ecosystem. Across all functional groups, there was evidence for nutritional control of resorption in this system, with both N and P resorption efficiencies decreasing as green leaf nutrient concentrations increased. However, specific patterns of resorption and the functional relationships between nutrient concentrations in green and senesced leaves varied by nutrient and plant functional group. Functional relationships between N concentrations in green and senesced leaves were indistinguishable between the dominant groups, suggesting that variation in N resorption efficiency was largely the result of inter-life form differences in green leaf N concentrations. In contrast, observed differences in P resorption efficiencies between life forms appear to be the result of both differences in green leaf P concentrations as well as inherent differences between life forms in the fraction of green leaf P resorbed from senescing leaves. Our results indicate that foliar N:P ratios are poor predictors of resorption efficiency in this ecosystem, in contrast to N and P resorption proficiencies, which are more responsive to foliar N:P ratios.

Contributions of woody and herbaceous vegetation to tropical savanna ecosystem productivity: a quasi-global estimate

To estimate the relative contributions of woody and herbaceous vegetation to savanna productivity, we measured the 13C/12C isotopic ratios of leaves from trees, shrubs, grasses and the surface soil carbon pool for 22 savannas in Australia, Brazil and Ghana covering the full savanna spectrum ranging from almost pure grassland to closed woodlands on all three continents. All trees and shrubs sampled were of the C3 pathway and all grasses of the C4 pathway with the exception of Echinolaena inflexa (Poir.) Chase, a common C3 grass of the Brazilian cerrado. By comparing the carbon isotopic compositions of the plant and carbon pools, a simple model relating soil delta 13C to the relative abundances of trees + shrubs (woody plants) and grasses was developed. The model suggests that the relative proportions of a savanna ecosystem's total foliar projected cover attributable to grasses versus woody plants is a simple and reliable index of the relative contributions of grasses and woody plants to savanna net productivity. Model calibrations against woody tree canopy cover made it possible to estimate the proportion of savanna productivity in the major regions of the world attributable to trees + shrubs and grasses from ground-based observational maps of savanna woodiness. Overall, it was estimated that 59% of the net primary productivity (Np) of tropical savannas is attributable to C4 grasses, but that this proportion varies significantly within and between regions. The C4 grasses make their greatest relative contribution to savanna Np in the Neotropics, whereas in African regions, a greater proportion of savanna Np is attributable to woody plants. The relative contribution of C4 grasses in Australian savannas is intermediate between those in the Neotropics and Africa. These differences can be broadly ascribed to large scale differences in soil fertility and rainfall.

Using Ninhydrin to Detect Gravesoil*

Some death scene investigations commence without knowledge of the location of the body and/or decomposition site. In these cases, it is necessary to locate the remains or the site where the body decomposed prior to movement. We hypothesized that the burial of a mammalian cadaver will result in the release of ninhydrin reactive nitrogen (NRN) into associated soil and that this reaction might have potential as a tool for the identification of clandestine graves. Juvenile rat (Rattus rattus) cadavers were buried in three contrasting soil types in Australian tropical savanna ecosystems and allowed to decompose over a period of 28 days. Soils were sequentially harvested and analyzed for NRN. Cadaver burial resulted in an approximate doubling (mean = 1.7 +/- 0.1) in the concentration of soil NRN. This reaction has great potential to be used as a presumptive test for gravesoil and this use might be greatly enhanced following more detailed research.

Stem and leaf hydraulics of congeneric tree species from adjacent tropical savanna and forest ecosystems

Leaf and stem functional traits related to plant water relations were studied for six congeneric species pairs, each composed of one tree species typical of savanna habitats and another typical of adjacent forest habitats, to determine whether there were intrinsic differences in plant hydraulics between these two functional types. Only individuals growing in savanna habitats were studied. Most stem traits, including wood density, the xylem water potential at 50% loss of hydraulic conductivity, sapwood area specific conductivity, and leaf area specific conductivity did not differ significantly between savanna and forest species. However, maximum leaf hydraulic conductance (K (leaf)) and leaf capacitance tended to be higher in savanna species. Predawn leaf water potential and leaf mass per area were also higher in savanna species in all congeneric pairs. Hydraulic vulnerability curves of stems and leaves indicated that leaves were more vulnerable to drought-induced cavitation than terminal branches regardless of genus. The midday K (leaf) values estimated from leaf vulnerability curves were very low implying that daily embolism repair may occur in leaves. An electric circuit analog model predicted that, compared to forest species, savanna species took longer for their leaf water potentials to drop from predawn values to values corresponding to 50% loss of K (leaf) or to the turgor loss points, suggesting that savanna species were more buffered from changes in leaf water potential. The results of this study suggest that the relative success of savanna over forest species in savanna is related in part to their ability to cope with drought, which is determined more by leaf than by stem hydraulic traits. Variation among genera accounted for a large proportion of the total variance in most traits, which indicates that, despite different selective pressures in savanna and forest habitats, phylogeny has a stronger effect than habitat in determining most hydraulic traits.

Autoclaving kills soil microbes yet soil enzymes remain active

Biocidal treatment of soil is used to remove or inhibit soil microbial activity, and thus provides insight into the relationship between soil biology and soil processes. Chemical (soil pH, phosphodiesterase, protease) and biological (substrate induced respiration) characteristics of three contrasting soils from tropical savanna ecosystems in north Queensland, Australia were measured in field fresh samples and following autoclaving (121 °C/103 kPa for 30 min on two consecutive days). Autoclaving treatment killed the active soil microbial biomass and significantly decreased protease activity (∼90%) in all three soils. Phosphodiesterase activity in kaolinitic soils also significantly decreased by 78% and 92%. However, autoclave treatment of smectitic soil only decreased phosphodiesterase activity by 4% only. This study demonstrates phosphodiesterase can remain stable in extreme conditions. This might be a characteristic vital to the cycling of phosphorus in shrink–swell clays in Australian tropical savanna ecosystems.

Productivity and carbon fluxes of tropical savannas

AbstractAim (1) To estimate the local and global magnitude of carbon fluxes between savanna and the atmosphere, and to suggest the significance of savannas in the global carbon cycle. (2) To suggest the extent to which protection of savannas could contribute to a global carbon sequestration initiative.Location Tropical savanna ecosystems in Africa, Australia, India and South America.Methods A literature search was carried out using the ISI Web of Knowledge, and a compilation of extra data was obtained from other literature, including national reports accessed through the personal collections of the authors. Savanna is here defined as any tropical ecosystem containing grasses, including woodland and grassland types. From these data it was possible to estimate the fluxes of carbon dioxide between the entire savanna biome on a global scale.Results Tropical savannas can be remarkably productive, with a net primary productivity that ranges from 1 to 12 t C ha−1 year−1. The lower values are found in the arid and semi‐arid savannas occurring in extensive regions of Africa, Australia and South America. The global average of the cases reviewed here was 7.2 t C ha−1 year−1. The carbon sequestration rate (net ecosystem productivity) may average 0.14 t C ha−1 year−1or 0.39 Gt C year−1. If savannas were to be protected from fire and grazing, most of them would accumulate substantial carbon and the sink would be larger. Savannas are under anthropogenic pressure, but this has been much less publicized than deforestation in the rain forest biome. The rate of loss is not well established, but may exceed 1% per year, approximately twice as fast as that of rain forests. Globally, this is likely to constitute a flux to the atmosphere that is at least as large as that arising from deforestation of the rain forest.Main conclusions The current rate of loss impacts appreciably on the global carbon balance. There is considerable scope for using many of the savannas as sites for carbon sequestration, by simply protecting them from burning and grazing, and permitting them to increase in stature and carbon content over periods of several decades.

Tree–grass coexistence in the Brazilian cerrado: demographic consequences of environmental instability

AbstractAim Tropical savanna ecosystems are uniquely characterized by the co‐dominance of both trees and grasses. An operational understanding of the ecological processes involved in maintaining this condition is essential for understanding both the functioning of savanna systems as well as their potential response to environmental change. A simple model is presented to explore the potential for a demographic mechanism of long‐term tree persistence and temporal physiognomic stability in the Brazilian cerrado.Location The model is developed based on data from the humid cerrado of Brazil.Methods In contrast to many existing models of tree–grass dynamics a model is presented which is based on data from the humid cerrado of Brazil, which is both qualitatively and quantitatively different from many of the more arid savannas of the palaeotropics. The model focuses on the dynamics of a synthetic tree population, with particular attention given to reproduction, seedling establishment and fire effects; with separate sub‐models for grass production, fire and rainfall.Results The model successfully predicts coexistence across the full range of observed vegetation physiognomies, but only under limited conditions. Under coexistence conditions, the dynamics of the tree population are characterized by long periods of gradual decline, punctuated by occasional bursts of growth. However, in agreement with earlier studies, the model consistently over‐predicts domination by the tree component. Fire is identified as an overriding factor in determining model behaviour, and the response of reproduction and sapling recruitment to variance in the frequency of fire ignition is identified to be of potential importance in the functioning of the Brazilian cerrado. The key dynamics of the model which promote tree–grass coexistence are consistent with a number of established determinants of ecological resilience in savanna systems.Main conclusions The model identifies the importance of the effective exploitation of rare opportunities for favourable recruitment (e.g. exclusion from fire) by the tree population, in promoting coexistence within a predominantly adverse environment. Support is provided for an alternative demographic mechanism of tree–grass coexistence in the cerrado (the storage effect), which is not based on the limiting assumption of niche partitioning through differences in rooting depth. The results are consistent with those presented by recent modelling work based on the more arid savannas of southern Africa. The model presented here differs in the emphasis given to particular environmental and life‐history attributes which are critical in determining the tree–grass balance, but provides further general support for the potential role of demographic mechanisms (such as the storage effect) in determining the structure of tropical savannas. Despite having clear limitations, models can serve as valuable heuristic tools to aid the integration and exploration of existing data sets as well as our present understanding of key ecological processes.

Stem and leaf gas exchange and their responses to fire in a north Australian tropical savanna

We measured stem CO2 efflux and leaf gas exchange in a tropical savanna ecosystem in northern Australia, and assessed the impact of fire on these processes. Gas exchange of mature leaves that flushed after a fire showed only slight differences from that of mature leaves on unburned trees. Expanding leaves typically showed net losses of CO2 to the atmosphere in both burned and unburned trees, even under saturating irradiance. Fire caused stem CO2 efflux to decline in overstory trees, when measured 8 weeks post-fire. This decline was thought to have resulted from reduced availability of C substrate for respiration, due to reduced canopy photosynthesis caused by leaf scorching, and to priority allocation of fixed C towards reconstruction of a new canopy. At the ecosystem scale, we estimated the annual above-ground woody-tissue CO2 efflux to be 275 g C m(-2) ground area year(-1) in a non-fire year, or approximately 13% of the annual gross primary production. We contrasted the canopy physiology of two co-dominant overstory tree species, one of which has a smooth bark on its branches capable of photosynthetic re-fixation (Eucalyptus miniata), and the other of which has a thick, rough bark incapable of re-fixation (Eucalyptus tetrodonta). Eucalyptus miniata supported a larger branch sapwood cross-sectional area in the crown per unit subtending leaf area, and had higher leaf stomatal conductance and photosynthesis than E. tetrodonta. Re-fixation by photosynthetic bark reduces the C cost of delivering water to evaporative sites in leaves, because it reduces the net C cost of constructing and maintaining sapwood. We suggest that re-fixation allowed leaves of E. miniata to photosynthesize at higher rates than those of E. tetrodonta, while the two invested similar amounts of C in the maintenance of branch sapwood.

Grass populations control nitrification in savanna soils

Summary Nitrification plays a key role in the functioning of many natural ecosystems. It is directly involved in plant nitrogen nutrition and soil N losses through leaching and denitrification. The control of this process by plants is poorly understood, although modifications of nitrification would allow plants to manipulate competition for N and induce changes in ecosystem N balance. In a wet tropical savanna ecosystem (Lamto, Côte d’Ivoire), the soil N cycle is characterized by distinct high‐ and low‐nitrification sites. Previous publications showed that nitrification was positively or negatively correlated with root densities of the dominant grass covering these sites. These contrasting sites were chosen to investigate the extent to which vegetation controls long‐term nitrification. In situ experimental plots were created where grass individuals originating from high‐ or low‐nitrifying soils were transplanted into both soils. Nitrifying enzyme activity (NEA) was measured up to 24 months after transplanting. Grasses from both sites significantly modified NEA up to rates similar to those at their respective control sites. The level of individual plant control (inhibition and stimulation) was correlated with grass biomass. The potential mechanisms of this control is discussed, along with its consequences for ecosystem N cycling (such as N losses), as the denitrifying enzyme activity (DEA) is much higher in the high‐nitrification site. Such results suggest that plant species can have important consequences for N cycling at the population level.

Historical C3-C4 vegetation pattern on forested mountain slopes: its implication for ecological rehabilitation of degraded highlands of Ethiopia by afforestation

In Ethiopia, plantation forestry for soil conservation and wood supply is mainly based on exotic tree species harvested at 12–25-y rotation age. To evaluate if these forests truly represent ecological rehabilitation of degraded areas through the build-up of soil organic matter before harvest, relative abundances of C3 vs. C4 carbon in soils under a 25-y-old forest on Mt. Yegof have been studied based on changes in soil δ13C values due to vegetation cover changes by afforestation. At Yegof, shrub and tree leaves had δ13C value of −28.7 ± 0.4‰. The grasses showed a value of −30.7 ± 0.6‰ at 2700–3000 m asl (typical C3) and of −13.6 ± 0.3‰ at 2520 m asl (typical C4). Soil δ13C values were −21.7 ± 0.9‰ in soil at 0–5 cm and −20.7 ± 0.6‰ in soil at 30–50 cm indicating a long history of C4-dominated grass or cropland over the elevations sampled. The shifts towards lower δ13C values in soil at 0–5 cm coincide with 25 y of C3 vegetation. Carbon content in topsoil was 3.4–9.2% and in tree leaves was 45–56%. However, an estimate of new C3 carbon (54%) in 0–5 cm soil was low as compared with tropical rain-forest and savanna ecosystems suggesting a slow rate of carbon turnover at Yegof. The results suggest that degraded sites at Yegof may need further duration of forest cover longer than current rotation times of the forests to improve soil conditions and restore soil organic matter and carbon, which have been lost during the past land use. It is encouraging from the perspectives of ecological rehabilitation and soil conservation that a shift in the dominance from C4 vegetation types of agricultural and grassland ecosystems to C3 forest ecosystems could be established in less than 25 y on these highly degraded slopes of Mt. Yegof.

Advantages and drawbacks of NOAA-AVHRR and SPOT-VGT for burnt area mapping in a tropical savanna ecosystem

In this paper we compare and contrast the performances of two low spatial resolution Earth observing systems, National Oceanographic and Atmospheric Administration ‐ advanced very high resolution radiometer (NOAA‐AVHRR) and Système pour l'Observation de la Terre ‐ VEGETATION (SPOT‐VGT), for burnt area mapping in a tropical savanna ecosystem of northern Australia. A multi-temporal composite image was derived from each of the two data sets acquired at the beginning of the 1999 fire season, and a set of rules for extracting burnt areas was compiled using a classification and regression trees (CART) methodology. In the case of the VGT data, the rules were used to segment the entire composite image; for the AVHRR data, a seed-growing approach was tested, with the aim of reducing overestimation of burnt areas. Since burnt areas are often confused with other land cover types, active fires were first identified (the seeds), and then the mapping of burnt areas was restricted to the neighbourhood of the pixels labelled as active fires. For the accuracy assessment, a reference map was derived by visual interpretation of two Landsat thematic mapper (TM) frames. First, the proportions of burnt areas were calculated for 14 × 14 km cells on a regular grid overlaid on AVHRR, VGT, and TM maps and the low-resolution areas were regressed against the TM estimates. Second, the spatial agreement between burnt area maps derived using high and low spatial resolution data was assessed using confusion matrices. We conclude that NOAA‐AVHRR and SPOT‐VGT show complementary characteristics. The better geometry of the VGT sensor, compared to the AVHRR, allows a higher accuracy in burnt area estimation, but the lack of a thermal channel on VGT is certainly a drawback for both the compositing and the burnt area mapping processes. Both systems tend to overestimate burnt areas when these areas are very fragmented but clearly underestimate when the burn scars are too small to be detected. Lastly, the seed-growing method is promising and could form the basis for a combined use of NOAA-AVHRR and SPOT-VGT systems.

Effect of fire and soil texture on soil carbon in a sub-humid savanna (Matopos, Zimbabwe)

We investigated the effects of changing fire regime on the stocks and isotopic composition of soil organic carbon (SOC) in a tropical savanna ecosystem at Matopos, Zimbabwe. Vegetation plots from both sandy and clay-rich soil types at this location have been subjected to fire frequencies ranging from annual burn to complete protection for the last 50 years. Gross variations in 0–5 cm SOC stocks and the δ 13C value of SOC were predominantly related to soil texture, with carbon densities at the sandy sites being consistently 35–50% lower than those at comparable clay sites. Average 0–5 cm carbon densities for all the burnt plots were approximately 100 mg/cm 2 and 50 mg/cm 2, at the clay site and the sandy site, respectively. In both cases, lower fire frequencies had resulted in a ∼10% increase, while higher fire frequencies had resulted in a ∼10% decrease from these average values. Plots from which fire had been excluded experienced a 40% to 50% increase in carbon stocks in the 0–5 cm interval, compared with the average for the burned plots. There was a linear relationship between carbon density and δ 13C value at both sandy and clay sites. This is controlled by the rate of delivery of C 3- and C 4-derived carbon to the SOC pool, by the differences in residence time for C 3- and C 4-derived carbon in the SOC pool (in turn controlled largely by fire frequency), and by soil texture. The distribution of carbon and 13C between size fractions is also controlled by soil texture and fire frequency. Increasing fire frequency results in a relative increase in fine particulate SOC and an increase in the δ 13C value of SOC in all size fractions. Soil texture, on the other hand, controls the magnitude of the increases in both the abundance and the δ 13C value of SOC in all size fractions.

Stable coexistence of contrasted nitrification statuses in a wet tropical savanna ecosystem

1. Wet tropical savannas are characterized by strong environmental constraints—particularly low soil nutrient availability—associated with high plant productivity. Nitrogen recycling, and especially nitrification, is supposed to be a strong determinant of the balance between conservation and loss of nutrients at the ecosystem level. Savanna facies dominated by the grass Hyparrhenia diplandra (Andropogoneae) are known to exhibit low levels of nitrification and thus avoid nitrate losses. 2. By comparing two sites in the Lamto area (Côte d’Ivoire, West Africa) with similar soil physico‐chemical characteristics and equally dominated by H. diplandra (80% of the grass cover), it was demonstrated that, within this facies, nitrification is highly heterogeneous, with a 240‐fold variation in potential nitrification within a specific site. 3. In order to test whether these differences can be considered as permanent in this ecosystem, nitrate reductase activities were compared on H. diplandra plantlets from the two sites, cultivated under identical conditions in the presence of nitrate. The leaves of plants originating from the high nitrification site were always able to reduce nitrate at a significantly higher rate than those from the low nitrification site. This observation indicates a long‐term adaptation of the plants and stable nitrification behaviour. 4. Lamto can thus be considered as a contrasted dual ecosystem relative to its nitrogen cycle. The two sites studied therefore constitute useful models to assess the determinism of nitrification in wet savannas and the role of this process on nitrogen retention in such ecosystems.