Is productivity of mesic savannas light limited or water limited? Results of a simulation study

A soil-plant-atmosphere model was used to estimate gross primary productivity (GPP) and evapotranspiration (ET) of a tropical savanna in Australia. This paper describes model modifications required to simulate the substantial C4 grass understory together with C3 trees. The model was further improved to include a seasonal distribution of leaf area and foliar nitrogen through 10 canopy layers. Model outputs were compared with a 5-year eddy covariance dataset. Adding the C4 photosynthesis component improved the model efficiency and root-mean-squared error (RMSE) for total ecosystem GPP by better emulating annual peaks and troughs in GPP across wet and dry seasons. The C4 photosynthesis component had minimal impact on modelled values of ET. Outputs of GPP from the modified model agreed well with measured values, explaining between 79% and 90% of the variance and having a low RMSE (0.003-0.281gCm-2day-1). Approximately, 40% of total annual GPP was contributed by C4 grasses. Total (trees and grasses) wet season GPP was approximately 75-80% of total annual GPP. Light-use efficiency (LUE) was largest for the wet season and smallest in the dry season and C4 LUE was larger than that of the trees. A sensitivity analysis of GPP revealed that daily GPP was most sensitive to changes in leaf area index (LAI) and foliar nitrogen (Nf) and relatively insensitive to changes in maximum carboxylation rate (Vcmax), maximum electron transport rate (Jmax) and minimum leaf water potential (ψmin). The modified model was also able to represent daily and seasonal patterns in ET, (explaining 68-81% of variance) with a low RMSE (0.038-0.19mmday-1). Current values of Nf, LAI and other parameters appear to be colimiting for maximizing GPP. By manipulating LAI and soil moisture content inputs, we show that modelled GPP is limited by light interception rather than water availability at this site. © 2011 Blackwell Publishing Ltd

Droughts and the ecological future of tropical savanna vegetation

1. Climate change is expected to lead to more frequent, intense and longer droughts in the future, with major implications for ecosystem processes and human livelihoods. The impacts of such droughts are already evident, with vegetation dieback reported from a range of ecosystems, including savannas, in recent years. 2. Most of our insights into the mechanisms governing vegetation drought responses have come from forests and temperate grasslands, while responses of savannas have received less attention. Because the two life forms that dominate savannas—C3 trees and C4 grasses—respond differently to the same environmental controls, savanna responses to droughts can differ from those of forests and grasslands. 3. Drought‐driven mortality of savanna vegetation is not readily predicted by just plant drought‐tolerance traits alone, but is the net outcome of multiple factors, including drought‐avoidance strategies, landscape and neighborhood context, and impacts of past and current stressors including fire, herbivory and inter‐life form competition. 4. Many savannas currently appear to have the capacity to recover from moderate to severe short‐term droughts, although recovery times can be substantial. Factors facilitating recovery include the resprouting ability of vegetation, enhanced flowering and seeding and post‐drought amelioration of herbivory and fire. Future increases in drought severity, length and frequency can interrupt recovery trajectories and lead to compositional shifts, and thus pose substantial threats, particularly to arid and semi‐arid savannas. 5. Synthesis. Our understanding of, and ability to predict, savanna drought responses is currently limited by availability of relevant data, and there is an urgent need for campaigns quantifying drought‐survival traits across diverse savannas. Importantly, these campaigns must move beyond reliance on a limited set of plant functional traits to identifying suites of physiological, morphological, anatomical and structural traits or “syndromes” that encapsulate both avoidance and tolerance strategies. There is also a critical need for a global network of long‐term savanna monitoring sites as these can provide key insights into factors influencing both resistance and resilience of different savannas to droughts. Such efforts, coupled with site‐specific rainfall manipulation experiments that characterize plant trait–drought response relationships, and modelling efforts, will enable a more comprehensive understanding of savanna drought responses

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. © Springer 2005

Modelling Seasonal and Inter-annual Variations in Carbon and Water Fluxes in an Arid-Zone Acacia Savanna Woodland, 1981–2012

© 2016, Springer Science+Business Media New York. Changes in climatic characteristics such as seasonal and inter-annual variability may affect ecosystem structure and function, hence alter carbon and water budgets of ecosystems. Studies of modelling combined with field experiments can provide essential information to investigate interactions between carbon and water cycles and climate. Here we present a first attempt to investigate the long-term climate controls on seasonal patterns and inter-annual variations in water and carbon exchanges in an arid-zone savanna-woodland ecosystem using a detailed mechanistic soil–plant–atmosphere model (SPA), driven by leaf area index (LAI) simulated by an ecohydrological model (WAVES) and observed climate data during 1981–2012. The SPA was tested against almost 3 years of eddy covariance flux measurements in terms of gross primary productivity (GPP) and evapotranspiration (ET). The model was able to explain 80 and 71% of the variability of observed daily GPP and ET, respectively. Long-term simulations showed that carbon accumulation rates and ET ranged from 20.6 g C m−2 mon−1 in the late dry season to 45.8 g C m−2 mon−1 in the late wet season, respectively, primarily driven by seasonal variations in LAI and soil moisture. Large climate variations resulted in large seasonal variation in ecosystem water-use efficiency (eWUE). Simulated annual GPP varied between 146.4 and 604.7 g C m−2 y−1. Variations in annual ET coincided with that of GPP, ranging from 110.2 to 625.8 mm y−1. Annual variations in GPP and ET were driven by the annual variations in precipitation and vapour pressure deficit (VPD) but not temperature. The linear coupling of simulated annual GPP and ET resulted in eWUE having relatively small year-to-year variation

Productivity and evapotranspiration of two contrasting semiarid ecosystems following the 2011 global carbon land sink anomaly

© 2016 Elsevier B.V. Global carbon balances are increasingly affected by large fluctuations in productivity occurring throughout semiarid regions. Recent analyses found a large C uptake anomaly in 2011 in arid and semiarid regions of the southern hemisphere. Consequently, we compared C and water fluxes of two distinct woody ecosystems (a Mulga (Acacia) woodland and a Corymbia savanna) between August 2012 and August 2014 in semiarid central Australia, demonstrating that the 2011 anomaly was short-lived in both ecosystems. The Mulga woodland was approximately C neutral but with periods of significant uptake within both years. The extreme drought tolerance of Acacia is presumed to have contributed to this. By contrast, the Corymbia savanna was a very large net C source (130 and 200gCm-2yr-1 in average and below average rainfall years, respectively), which is likely to have been a consequence of the degradation of standing, senescent biomass that was a legacy of high productivity during the 2011 anomaly. The magnitude and temporal patterns in ecosystem water-use efficiencies (WUE), derived from eddy covariance data, differed across the two sites, which may reflect differences in the relative contributions of respiration to net C fluxes across the two ecosystems. In contrast, differences in leaf-scale measures of WUE, derived from 13C stable isotope analyses, were apparent at small spatial scales and may reflect the different rooting strategies of Corymbia and Acacia trees within the Corymbia savanna. Restrictions on root growth and infiltration by a siliceous hardpan located below Acacia, whether in the Mulga woodland or in small Mulga patches of the Corymbia savanna, impedes drainage of water to depth, thereby producing a reservoir for soil moisture storage under Acacia while acting as a barrier to access of groundwater by Corymbia trees in Mulga patches, but not in the open Corymbia savanna

Deforestation for Pasture Development – Has It Been Worth It?

Differing scenarios leading to deforestation for pasture development in savanna (woodland) and closed forest communities in the tropics – sub-tropics are compared and contrasted. Australian and Brazilian examples are highlighted. No simple answer is given to the question of whether deforestation for pasture development has been worth it, since both commercial and non-commercial values have equal validity and need to be taken into account. These issues are addressed in the context of land assigned by governments for agricultural purposes. It is concluded that technology and ecological understanding are now available to maintain sustainable production from converted forest systems. However emphasis should be on delivering this within the framework of existing deforested areas – rather than in expanding the area of forest conversion

How energy and water availability constrain vegetation water-use along the North Australian Tropical Transect

© 2016, Gorgan Univ Agricultural Sciences and Natural Resources. All rights reserved. Energy and water availability were identified as the first order controls of evapotranspiration (ET) in ecohyrodrology. With a ~1,000 km precipitation gradient and distinct wet-dry climate, the North Australian Tropical Transect (NATT) was well suited for evaluating how energy and water availabilities constrain water use by vegetation, but has not been done yet. In this study, we addressed this question using Budyko framework that quantifies the evapotranspiration as a function of energy-limited rate and precipitation. Path analysis was adopted to evaluate the dependencies of water and carbon fluxes on ecohydrological variables. Results showed that the major drivers of water and carbon fluxes varied between wet and dry savannas: down-welling solar radiation was the primary driver of the wet season ET in mesic savanna ecosystems, while soil water availability was the primary driver in inland dryland ecosystems. Vegetation can significantly regulate water and carbon fluxes of savanna ecosystems, as supported by the strong link of LAI with ET and GPP from path analysis. Vegetation structure (i.e. the tree:grass ratio) at each site can regulate the impact of climatic constraint on ET and GPP. Sites with a low tree:grass ratio had ET and GPP that exceeded sites with high a tree:grass ratio when the grassy understory was active. Identifying the relative importance of these climate drivers and vegetation structure on seasonal patterns of water use by these ecosystems will help us decide our priorities when improving the estimates of ET and GPP

Coupling carbon allocation with leaf and root phenology predicts tree-grass partitioning along a savanna rainfall gradient

© Author(s) 2016. The relative complexity of the mechanisms underlying savanna ecosystem dynamics, in comparison to other biomes such as temperate and tropical forests, challenges the representation of such dynamics in ecosystem and Earth system models. A realistic representation of processes governing carbon allocation and phenology for the two defining elements of savanna vegetation (namely trees and grasses) may be a key to understanding variations in tree-grass partitioning in time and space across the savanna biome worldwide. Here we present a new approach for modelling coupled phenology and carbon allocation, applied to competing tree and grass plant functional types. The approach accounts for a temporal shift between assimilation and growth, mediated by a labile carbohydrate store. This is combined with a method to maximize long-term net primary production (NPP) by optimally partitioning plant growth between fine roots and (leaves + stem). The computational efficiency of the analytic method used here allows it to be uniquely and readily applied at regional scale, as required, for example, within the framework of a global biogeochemical model. We demonstrate the approach by encoding it in a new simple carbon-water cycle model that we call HAVANA (Hydrology and Vegetation-dynamics Algorithm for Northern Australia), coupled to the existing POP (Population Orders Physiology) model for tree demography and disturbance-mediated heterogeneity. HAVANA-POP is calibrated using monthly remotely sensed fraction of absorbed photosynthetically active radiation (fPAR) and eddy-covariance-based estimates of carbon and water fluxes at five tower sites along the North Australian Tropical Transect (NATT), which is characterized by large gradients in rainfall and wildfire disturbance. The calibrated model replicates observed gradients of fPAR, tree leaf area index, basal area, and foliage projective cover along the NATT. The model behaviour emerges from complex feedbacks between the plant physiology and vegetation dynamics, mediated by shifting above- versus below-ground resources, and not from imposed hypotheses about the controls on tree-grass co-existence. Results support the hypothesis that resource limitation is a stronger determinant of tree cover than disturbance in Australian savannas

Coupling carbon allocation with leaf and root phenology predicts tree-grass partitioning along a savanna rainfall gradient

The relative complexity of the mechanisms underlying savanna ecosystem dynamics, in comparison to other biomes such as temperate and tropical forests, challenges the representation of such dynamics in ecosystem and Earth system models. A realistic representation of processes governing carbon allocation and phenology for the two defining elements of savanna vegetation (namely trees and grasses) may be a key to understanding variations in tree–grass partitioning in time and space across the savanna biome worldwide. Here we present a new approach for modelling coupled phenology and carbon allocation, applied to competing tree and grass plant functional types. The approach accounts for a temporal shift between assimilation and growth, mediated by a labile carbohydrate store. This is combined with a method to maximize long-term net primary production (NPP) by optimally partitioning plant growth between fine roots and (leaves + stem). The computational efficiency of the analytic method used here allows it to be uniquely and readily applied at regional scale, as required, for example, within the framework of a global biogeochemical model. We demonstrate the approach by encoding it in a new simple carbon–water cycle model that we call HAVANA (Hydrology and Vegetation-dynamics Algorithm for Northern Australia), coupled to the existing POP (Population Orders Physiology) model for tree demography and disturbance-mediated heterogeneity. HAVANA-POP is calibrated using monthly remotely sensed fraction of absorbed photosynthetically active radiation (fPAR) and eddy-covariance-based estimates of carbon and water fluxes at five tower sites along the North Australian Tropical Transect (NATT), which is characterized by large gradients in rainfall and wildfire disturbance. The calibrated model replicates observed gradients of fPAR, tree leaf area index, basal area, and foliage projective cover along the NATT. The model behaviour emerges from complex feedbacks between the plant physiology and vegetation dynamics, mediated by shifting above- versus below-ground resources, and not from imposed hypotheses about the controls on tree–grass co-existence. Results support the hypothesis that resource limitation is a stronger determinant of tree cover than disturbance in Australian savannas.The contributions of V. Haverd and P. Briggs were made possible by the support of the Australian Climate Change Science Program. B. Smith acknowledges funding as an OCE Distinguished Visiting Scientist to the CSIRO Oceans & Atmosphere Flagship, Canberr