Fire in Australian savannas: from leaf to landscape

Savanna ecosystems comprise 22% of the global terrestrial surface and 25% of Australia (almost 1.9 million km²) and provide significant ecosystem services through carbon and water cycles and the maintenance of biodiversity. The current structure, composition and distribution of Australian savannas have coevolved with fire, yet remain driven by the dynamic constraints of their bioclimatic niche. Fire in Australian savannas influences both the biophysical and biogeochemical processes at multiple scales from leaf to landscape. Here, we present the latest emission estimates from Australian savanna biomass burning and their contribution to global greenhouse gas budgets. We then review our understanding of the impacts of fire on ecosystem function and local surface water and heat balances, which in turn influence regional climate. We show how savanna fires are coupled to the global climate through the carbon cycle and fire regimes. We present new research that climate change is likely to alter the structure and function of savannas through shifts in moisture availability and increases in atmospheric carbon dioxide, in turn altering fire regimes with further feedbacks to climate. We explore opportunities to reduce net greenhouse gas emissions from savanna ecosystems through changes in savanna fire management.     

Soil organic carbon content at a range of north Australian tropical savannas with contrasting site histories

Soils play an important role in the global carbon cycle, and can be major source or sink of CO₂ depending upon land use, vegetation type and soil management practices. Natural and human impact on soil carbon concentration and storage is poorly understood in native north Australian savanna, yet this represents the largest carbon store in the ecosystem. To gain understanding of possible management impacts on this carbon pool, soil organic carbon (SOC) of the top 1m of red earth sands and sandy loams common in the region was sampled at 5 sites with different vegetation cover and site history (fire regime and tree removal). SOC was high when compared to other published values for savannas and was more comparable with dry-deciduous tropical forests. Sites sampled in this study represent high rainfall savannas of northern Australia (> 1700 mm annual rainfall) that feature frequent burning (2 in 3 years or more frequent) and a cycle of annual re-growth of tall C₄ grasses that dominate the savanna understorey. These factors may be responsible for the higher than expected SOC levels of the surface soils, despite high respiration rates. Medium term fire exclusion (15–20 years) at one of the sampled sites (Wildlife Park) dramatically reduced the grassy biomass of the understorey. This site had lower SOC levels when compared to the grass dominated and frequently burnt sites, which may be due to a reduction in detrital input to surface (0–30 cm) soil carbon pools. Exclusion of trees also had a significant impact on both the total amount and distribution of soil organic carbon, with tree removal reducing observed SOC at depth (100 cm). Soil carbon content was higher in the wet season than that in the dry season, but this difference was not statistically significant. Our results indicated that annual cycle of grass growth and wildfire resulted in small carbon accumulation in the upper region of the soil, and removal of woody plants resulted in significant carbon losses to recalcitrant, deep soil horizons greater than 80 cm depth.     

Seasonal changes in hydraulic conductance, xylem embolism and leaf area in Eucalyptus tetrodonta and Eucalyptus miniata saplings in a north Australian savanna

Eucalypt saplings in north Australian savannas commonly die back, sometimes to ground level, during the 5 months of the long dry season. Water potentials are lower in saplings than large trees during the dry season, and we hypothesized that low water potentials may lead to high levels of xylem embolism and consequent death of branches and whole shoots. As the dry season progressed, hydraulic conductance of terminal branches decreased by 50% in Eucalyptus tetrodonta but not in Eucalyptus miniata saplings. Hydraulic conductance per leaf area decreased seasonally by 34% in E. tetrodonta branches. These decreases may be associated with the loss of leaves recorded from E. tetrodonta but not E. miniata branches. We modelled the effect of sequential loss of parallel resistors, representing petioles on a branch. This showed there is a non-linear decrease in flow as basal resistors are lost, which can lead to a decrease in mean flow per resistor due to increased mean path-length. Thus the observed loss of basal leaves, together with xylem embolism, probably contributed to the seasonal loss of hydraulic conductance in E. tetrodonta saplings. Loss of hydraulic conductance due to xylem embolism was generally low ( < 15%) in both species, and the seasonal increase in embolism could not fully account for the decline in hydraulic conductance of E. tetrodonta branches. There was little evidence that branch and shoot death was caused by these levels of embolism. Developing an embolism vulnerability curve for species with long vessels is problematic and this issue is discussed.     

Episodic seedling growth in Allosyncarpia ternata,a lignotuberous,monsoon rainforest tree in northern Australia

On the western Arnhem Land Plateau, Northern Territory, Australia, seedlings of the canopy tree Allosyncarpia ternata S.T. Blake typically spend many years (perhaps decades) as small (<1 m), multistemmed plants on the forest floor. In this establishment phase, long periods of apparent inactivity are interrupted by episodes of rapid growth. This paper describes a 5‐year field‐monitoring program to examine the pattern of seedling growth and survival in allosyncarpia forest, and field and shadehouse measurements of lignotuber size. Individual seedlings may produce, each wet season, a number of fast‐growing stems, which then die back in the following dry season. As a result, mean annual above‐ground growth during this life stage is negligible. With each wet season, however, the seedling extends its below ground parts – a large lignotuber and a deep root system. After a number of years, when the lignotuber has grown large enough to sustain massive shoot growth, when a suitable light gap becomes available, and presumably when roots reach reliable dry‐season water supplies, the seedling grows rapidly. Thus, the shortage of saplings in allosyncarpia forest is due to the short time that individual plants spend at that particular growth‐stage, rather than to any dysfunction in recruitment.     

The response of sap flow to pulses of rain in a temperate Australian woodland

In water-limited systems, pulses of rainfall can trigger a cascade of plant physiological responses. However, the timing and size of the physiological response can vary depending on plant and environmental characteristics, such as rooting depth, plant size, rainfall amount, or antecedent soil moisture. We investigated the influence of pulses of rainfall on the response of sap flow of two dominant evergreen tree species, Eucalyptus crebra (a broadleaf) and Callitris glaucophylla (a needle leaved tree), in a remnant open woodland in eastern Australia. Sap flow data were collected using heat-pulse sensors installed in six trees of each species over a 2 year period which encompassed the tail-end of a widespread drought. Our objectives were to estimate the magnitude that a rainfall pulse had to exceed to increase tree water use (i.e., define the threshold response), and to determine how tree and environmental factors influenced the increase in tree water use following a rainfall pulse. We used data filtering techniques to isolate rainfall pulses, and analysed the resulting data with multivariate statistical analysis. We found that rainfall pulses less than 20 mm did not significantly increase tree water use (P > 0.05). Using partial regression analysis to hold all other variables constant, we determined that the size of the rain event (P < 0.05, R ² = 0.59), antecedent soil moisture (P < 0.05, R ² = 0.29), and tree size (DBH, cm, P < 0.05, R ² = 0.15), all significantly affected the response to rainfall. Our results suggest that the conceptual Threshold-Delay model describing physiological responses to rainfall pulses could be modified to include these factors. We further conclude that modelling of stand water use over an annual cycle could be improved by incorporating the T-D behaviour of tree transpiration. Responsible Editor: Stephen S.O. Burgess     

Monsoonal influences on evapotranspiration of savanna vegetation of northern Australia

Data from savannas of northern Australia are presented for net radiation, latent and sensible heat, ecosystem surface conductance (G_s) and stand water use for sites covering a latitudinal range of 5° or 700 km. Measurements were made at three locations of increasing distance from the northern coastline and represent high- (1,750 mm), medium- (890 mm) and low- (520 mm) rainfall sites. This rainfall gradient arises from the weakened monsoonal influence with distance inland. Data were coupled to seasonal estimates of leaf area index (LAI) for the tree and understorey strata. All parameters were measured at the seasonal extremes of late wet and dry seasons. During the wet season, daily rates of evapotranspiration were 3.1-3.6 mm day^-1 and were similar for all sites along the rainfall gradient and did not reflect site differences in annual rainfall. During the dry season, site differences were very apparent with evapotranspiration 2-18 times lower than wet season rates, the seasonal differences increasing with distance from coast and reduced annual rainfall. Due to low overstorey LAI, more than 80% of water vapour flux was attributed to the understorey. Seasonal differences in evapotranspiration were mostly due to reductions in understorey leaf area during the dry season. Water use of individual trees did not differ between the wet and dry seasons at any of the sites and stand water use was a simple function of tree density. G_s declined markedly during the dry season at all sites, and we conclude that the savanna water (and carbon) balance is largely determined by G_s and its response to atmospheric and soil water content and by seasonal adjustments to canopy leaf area.     

A Cost-Benefit Analysis of Leaves of Eight Australian Savanna Tree Species of Differing Leaf Life-Span

Cost-benefit analysis of foliar construction and maintenance costs and of carbon assimilation of leaves of differing life-span were conducted using two evergreen, three semi-deciduous, and three deciduous tree species of savannas of north Australia. Rates of radiant-energy-saturated CO₂ assimilation (P _max) and dark respiration were measured and leaves were analysed for total nitrogen, fat, and ash concentrations, and for heat of combustion. Specific leaf area, and leaf N and ash contents were significantly lower in longer-lived leaves (evergreen) than shorter-lived leaves (deciduous) species. Leaves of evergreen species also had significantly higher heat of combustion and lower crude fat content than leaves of deciduous species. On a leaf area basis, P _max was highest in leaves of evergreen species, but on a leaf dry mass basis it was highest in leaves of deciduous species. P _max and total Kieldahl N content were linearly correlated across all eight species, and foliar N content was higher in leaves of deciduous than evergreen species. Leaf construction cost was significantly higher and maintenance costs were lower for leaves of evergreen than deciduous species. Maintenance and construction costs were linearly related to each other across all species. Leaves of evergreen species had a higher cost-benefit ratio compared to leaves of deciduous species but with longer lived leaves, the payback interval was longer in evergreen than deciduous species. These results support the hypotheses that longer lived leaves are more expensive to construct than short-lived leaves, and that a higher investment of N into short-lived leaves occurs which supports a higher P _max over a shorter payback interval. This revised version was published online in June 2006 with corrections to the Cover Date.     

A Cost-Benefit Analysis of Leaves of Eight Australian Savanna Tree Species of Differing Leaf Life-Span

Cost-benefit analysis of foliar construction and maintenance costs and of carbon assimilation of leaves of differing life-span were conducted using two evergreen, three semi-deciduous, and three deciduous tree species of savannas of north Australia. Rates of radiant-energy-saturated CO₂ assimilation (P _max) and dark respiration were measured and leaves were analysed for total nitrogen, fat, and ash concentrations, and for heat of combustion. Specific leaf area, and leaf N and ash contents were significantly lower in longer-lived leaves (evergreen) than shorter-lived leaves (deciduous) species. Leaves of evergreen species also had significantly higher heat of combustion and lower crude fat content than leaves of deciduous species. On a leaf area basis, P _max was highest in leaves of evergreen species, but on a leaf dry mass basis it was highest in leaves of deciduous species. P _max and total Kieldahl N content were linearly correlated across all eight species, and foliar N content was higher in leaves of deciduous than evergreen species. Leaf construction cost was significantly higher and maintenance costs were lower for leaves of evergreen than deciduous species. Maintenance and construction costs were linearly related to each other across all species. Leaves of evergreen species had a higher cost-benefit ratio compared to leaves of deciduous species but with longer lived leaves, the payback interval was longer in evergreen than deciduous species. These results support the hypotheses that longer lived leaves are more expensive to construct than short-lived leaves, and that a higher investment of N into short-lived leaves occurs which supports a higher P _max over a shorter payback interval.