Lindsay B. Hutley

Savanna vegetation-fire-climate relationships differ among continents.

Surveying Savannas Savannas are structurally similar across the three major continents where they occur, leading to the assumption that the factors controlling vegetation structure and function are broadly similar, too. Lehmann et al. (p. 548) report the results of an extensive analysis of ground-based tree abundance in savannas, sampled at more than 2000 sites in Africa, Australia, and South America. All savannas, independent of region, shared a common functional property in the way that moisture and fire regulated tree abundance. However, despite qualitative similarity in the moisture–fire–tree-biomass relationships among continents, key quantitative differences exist among the three regions, presumably as a result of unique evolutionary histories and climatic domains. Evolution cannot be overlooked when aiming to predict the potential global impacts on savanna dynamics in a warming world. Ecologists have long sought to understand the factors controlling the structure of savanna vegetation. Using data from 2154 sites in savannas across Africa, Australia, and South America, we found that increasing moisture availability drives increases in fire and tree basal area, whereas fire reduces tree basal area. However, among continents, the magnitude of these effects varied substantially, so that a single model cannot adequately represent savanna woody biomass across these regions. Historical and environmental differences drive the regional variation in the functional relationships between woody vegetation, fire, and climate. These same differences will determine the regional responses of vegetation to future climates, with implications for global carbon stocks.

Fire impacts on surface heat, moisture and carbon fluxes from a tropical savanna in northern Australia

Savannas form a large fraction of the total tropical vegetation and are extremely fire prone. We measured radiative, energy and carbon exchanges over unburned and burned (both before and after low and moderate intensity fires) open forest savanna at Howard Springs, Darwin, Australia. Fire affected the radiative balance immediately following fire through the consumption of the grass-dominated understorey and blackening of the surface. Albedo was halved following fire of both intensities (from 0.12 to 0.07 and from 0.11 to 0.06 for the moderate and low intensity sites, respectively), but the recovery of albedo was dependent on the initial fire intensity. The low intensity fire caused little canopy damage with little impact on the surface energy balance and only a slight increase in Bowen ratio. However the moderate fire resulted in a comprehensive canopy scorch and almost complete leaf drop in the weeks following fire. The shutdown of most leaves within the canopy reduced transpiration and altered energy partitioning. Leaf death and shedding also resulted in a cessation of ecosystem carbon uptake and the savanna turned from a sink to a source of carbon to the atmosphere because of the continued ecosystem respiration. Post-fire, the Bowen ratio increased greatly due to large increases in sensible heat fluxes. These changes in surface energy exchange following fire, when applied at the landscape scale, may have impacts on climate through local changes in circulation patterns and changes in regional heating, precipitation and monsoon circulation.

Composition, leaf area index and standing biomass of eucalypt open forests near Darwin in the Northern Territory, Australia

Savanna communities dominate the wet–dry tropical regions of the world and are an important community type in monsoonal northern Australia. As such they have a significant impact on the water and carbon balance of this region. Above the 1200-mm isohyet, savanna’s are dominated by Eucalyptus miniata–E. tetrodonta open forests. We have described in detail the composition and structure as well as seasonal patterns of leaf area index and above-ground biomass in the E. miniata–E. tetrodonta open forests of the Gunn Point region near Darwin in the Northern Territory of Australia. In all, 29 tree species from four phenological guilds were recorded in these forests. Stand structure suggests that the forests were still recovering from the impacts of cyclone Tracy and subsequent frequent fires. Eucalyptus miniata and E. tetrodonta were significant contributors to overstorey leaf area index and standing biomass (>70%), and both leaf area index and biomass were strongly correlated to basal area. Leaf area index was at a maximum (about 1.0) at the end of the wet season and declined over the dry season by about 30–40%. There were proportionally greater changes in the understorey reflecting the greater contribution of deciduous and semi-deciduous species in this strata. Standing biomass was about 55 t ha –1 . Detailed descriptions of leaf area index and biomass are important inputs into the development of a water and carbon balance for the savanna’s of northern Australia.

Assessing the carbon sequestration potential of mesic savannas in the Northern Territory, Australia: approaches, uncertainties and potential impacts of fire

Tropical savannas cover a quarter of the Australian landmass and the biome represents a significant potential carbon sink. However, these savannas are subject to frequent and extensive fire. Fire regimes are likely to affect the productivity and carbon sequestration potential of savannas, through effects on both biomass and carbon emissions. The carbon sequestration potential has been estimated for some savanna sites by quantifying carbon storage in biomass and soil pools, and the fluxes to these pools. Using different techniques, previous work in these savannas has indicated that net ecosystem productivity [NEP, net primary productivity (NPP) less heterotrophic respiration] was about -3 t C ha-1 y-1 (i.e. a carbon sink). However, the impacts of fire were not accounted for in these calculations. Estimates of NEP have been combined with remotely-sensed estimates of area burnt and associated emissions for an extensive area of mesic savanna in Arnhem Land, NT, Australia. Combining NEP estimates with precise fire data provides an estimate of net biome productivity (NBP), a production index that includes carbon loss through disturbance (fire), and is thus a more realistic indicator of sequestration rate from this biome. This preliminary analysis suggests that NBP is approximately -1 t C ha-1 y-1 (i.e. a carbon sink). A reduction in the annual area burnt is likely to increase the sink size. Uncertainties surrounding these estimates of NBP and the implications of these uncertainties for land management in these extensive landscapes are discussed.

The utility of the eddy covariance techniques as a tool in carbon accounting: tropical savanna as a case study

Global concern over rising atmospheric CO2 concentrations has led to a proliferation of studies of carbon cycling in terrestrial ecosystems. Associated with this has been widespread adoption of the eddy covariance method to provide direct estimates of mass and energy exchange between vegetation surfaces and the atmosphere. With the eddy covariance method, fast-response instruments (10–20Hz) are typically mounted above plant canopies and the fluxes are calculated by correlating turbulent fluctuations in vertical velocity with fluctuations in various scalars such as CO2, water vapour and temperature. These techniques allow the direct and non-destructive measurement of the net exchange of CO2 owing to uptake via photosynthesis and loss owing to respiration, evapotranspiration and sensible heat. Eddy covariance measurements have a high temporal resolution, with fluxes typically calculated at 30-min intervals and can provide daily, monthly or annual estimates of carbon uptake or loss from ecosystems. Such measurements provide a bridge between ‘bottom-up’ (e.g. leaf, soil and whole plant measures of carbon fluxes) and ‘top-down’ approaches (e.g. satellite remote sensing, air sampling networks, inverse numerical methods) to understanding carbon cycling. Eddy covariance data also provide key measurements to calibrate and validate canopy- and regional-scale carbon balance models. Limitations of the method include high establishment costs, site requirements of flat and relatively uniform vegetation and problems estimating fluxes accurately at low wind speeds. Advantages include spatial averaging over 10–100ha and near-continuous measurements. The utility of the method is illustrated in current flux studies at ideal sites in northern Australia. Flux measurements spanning 3years have been made at a mesic savanna site (Howard Springs, Northern Territory) and semi-arid savanna (Virginia Park, northern Queensland). Patterns of CO2 and water vapour exchange at diurnal, seasonal and annual scales are detailed. Carbon dynamics at these sites are significantly different and reflect differences in climate and land management (impacts of frequent fire and grazing). Such studies illustrate the utility of the eddy covariance method and its potential to provide accurate data for carbon accounting purposes. If full carbon accounting is implemented, for ideal sites, the eddy covariance method provides annual estimates of carbon sink strength accurate to within 10%. The impact of land-use change on carbon sink strength could be monitored on a long-term basis and provide a valuable validation tool if carbon trading schemes were implemented.

Root biomass and root fractal analyses of an open Eucalyptus forest in a savanna of north Australia

Below-ground biomass of a Eucalyptus savanna forest was estimated following trenching to depths of 2 m around 16 mature trees in a tropical savanna of north Australia. Correlations among below-ground and various components of above-ground biomass were also investigated. In addition, root morphology was investigated by fractal analyses and a determination of an index of shallow-rootedness was undertaken. Total root biomass was 38.4 t ha–1, including 1 t ha–1 of fine roots. About 77–90p of total root biomass was found in the upper 0.5 m of soil. While fine-root biomass density was approximately constant (0.1 kg m–3) in the top soil, irrespective of distance from a tree stem, coarse-root biomass showed large variation with distance from the tree stem. Significant positive correlations among total root biomass, total above-ground biomass, diameter at breast height, leaf biomass and leaf area were obtained. It is likely that total root biomass can be reasonably accurately estimated from aboveground biomass and fine-root biomass from tree leaf area. We present equations that allow the prediction of belowground biomass from above-ground measures of tree size. Root morphology of two evergreen and two deciduous species was compared by the use of three parameters. These were the fractal dimension (d), which describes root system complexity; a proportionality factor (α), which is the ratio of the cross-sectional area before and after branching; and two indices of shallow-rootedness (ISR). Roots were found to be amenable to fractal analyses. The proportionality factor was independent of root diameter (Dr) at any branching level in all tree species examined, indicating that branching patterns were similar across all root sizes. The fractal dimension (d) ranged from 1.15 to 1.36, indicating a relatively simple root structure. Mean d was significantly different between E. tetrodonta (evergreen) and T. ferdinandiana (deciduous); however, no significant differences were found among other pairs of species. Terminalia ferdinandiana had the highest ISR, while Planchonia careya (deciduous) had the lowest. In addition, differences in ISR between P. careya and the other three species were significant, but not significant among E. miniata, E. tetrodonta and T. ferdinandiana. There were clear relationships among above-ground tree stem diameter at breast height, stem base diameter, and horizontal and vertical proximal root diameter. By the use of mean values of and stem diameter, we estimated the total crosssectional area of root and root diameter-class distribution for each species studied.

Control of atmospheric particles on diffuse radiation and terrestrial plant productivity: A review

Terrestrial plant productivity tends to increase under increasing but non-saturating photosynthetically active solar radiation when water, temperature and nutrients are not limiting. However, studies have shown that photosynthesis can also be higher under enhanced diffuse light despite a decrease in total irradiance. Clouds and atmospheric aerosols are two important variables that determine the total and proportion of diffuse light reaching the surface and thereby the rate of photosynthesis and carbon accumulation in plants. In addition to these factors, the response of plants to diffuse radiation is also dependant on plant characteristics such as functional types, leaf physiology, leaf area, leaf inclination, canopy structure and shape (i.e. clumping). Local environmental conditions (i.e. temperature, soil moisture, vapour pressure deficit, etc.) then modulate these plant responses. Changes in solar radiation as a consequence of clouds and aerosols thus can modify the carbon balance of terrestrial ecosystems. Therefore, understanding the role of solar radiation in terrestrial carbon processes has become one of the goals in terrestrial carbon cycle studies. It can help to identify the control and mechanisms of carbon processes and determines the geographical and temporal distribution of the major pools and fluxes in the global carbon cycle. Here we review the role of clouds and aerosols in partitioning solar radiation and their interactions with carbon processes of terrestrial plants. We also focus our review on vegetation characteristics that control the impact of radiation partitioning on vegetation carbon processes and the role of modelling approach to study this impact. We identify gaps in this field of research and further propose recommendations to bridge the gap.

The estimation of carbon budgets of frequently burnt tree stands in savannas of northern Australia, using allometric analysis and isotopic discrimination

The stock, rates of sequestration and allocation of carbon were estimated for trees in 14 0.1-ha plots at Kapalga in Kakadu National Park, Northern Territory, using new allometric relationships of carbon stock to stem cross-sectional area and measured growth rates of trees. Carbon stocks of trees ranged from 12 to 58 t ha–1, with sequestration representing ~9% of the total stocks. More than half of the sequestered carbon is allocated to leaves and twigs and ~20% to wood. Only ~25% is retained in the live trees with leaf and twig fall accounting for 80%–84% of the total transfers to the environment. An alternative method of calculating sequestration rates from consideration of water use and carbon-isotope discrimination data had a close to 1 : 1 match with estimates from allometric relationships. We developed and applied algorithms to predict the impacts of fire on carbon stocks of live trees. This showed that the reduction in live carbon stocks caused by single fires increased with increasing intensity, but the impact was highly dependent on the tree stand structure.

Seasonal patterns of soil carbon dioxide efflux from a wet-dry tropical savanna of northern Australia

Soil CO2 efflux rates were measured in a eucalypt open forest in a tropical savanna of northern Australia, with a portable closed chamber and CO2 gas analyser. Both abiotic (soil temperature and water content) and biotic (litterfall and fine-root growth) factors that may influence soil CO2 efflux were examined. Daytime rates of soil CO2 efflux rate were consistently higher than nocturnal values. Maximal rates occurred during late afternoons when soil temperatures were also maximal and minimum values were recorded during the early morning (0400-0800 hours). Average soil CO2 efflux was 5.37 mol m -2 s -1 (range 3.5-6.7 mol m -2 s -1 ) during the wet season and declined to 2.20 mol m -2 s -1 (range 1.2-3.6 mol m -2 s -1 ) during the dry season. The amount of carbon released from soil was 14.3 t ha -1 year -1 , with approximately 70% released during the wet season and 30% during the dry season. The rate of efflux was correlated with soil moisture content and soil temperature only during the wet season, when root growth and respiration were high. During the dry season there was no correlation with soil temperature. These results are discussed in relation to the carbon balance of tropical savannas.

Allometry for estimating aboveground tree biomass in tropical and subtropical eucalypt woodlands: towards general predictive equations

A fundamental tool in carbon accounting is tree-based allometry, whereby easily measured variables can be used to estimate aboveground biomass (AGB). To explore the potential of general allometry we combined raw datasets from 14 different woodland species, mainly eucalypts, from 11 sites across the Northern Territory, Queensland and New South Wales. Access to the raw data allowed two predictor variables, tree diameter (at 1.3-m height; D) and tree height (H), to be used singly or in various combinations to produce eight candidate models. Following natural log (ln) transformation, the data, consisting of 220 individual trees, were re-analysed in two steps: first as 20 species–site-specific AGB equations and, second, as a single general AGB equation. For each of the eight models, a comparison of the species–site-specific with the general equations was made with the Akaike information criterion (AIC). Further model evaluation was undertaken by a leave-one-out cross-validation technique. For each of the model forms, the species–site-specific equations performed better than the general equation. However, the best performing general equation, ln(AGB) = –2.0596 + 2.1561 ln(D) + 0.1362 (ln(H))2, was only marginally inferior to the species–site-specific equations. For the best general equation, back-transformed predicted v. observed values (on a linear scale) were highly concordant, with a slope of 0.99. The only major deviation from this relationship was due to seven large, hollow trees (more than 35% loss of cross-sectional stem area at 1.3 m) at a single species–site combination. Our best-performing general model exhibited remarkable stability across species and sites, when compared with the species–site equations. We conclude that there is encouraging evidence that general predictive equations can be developed across sites and species for Australia’s woodlands. This simplifies the conversion of long-term inventory measurements into AGB estimates and allows more resources to be focused on the extension of such inventories.