When is a ‘forest’ a savanna,and why does it matter?

Savannas are defined based on vegetation structure, the central concept being a discontinuous tree cover in a continuous grass understorey. However, at the high‐rainfall end of the tropical savanna biome, where heavily wooded mesic savannas begin to structurally resemble forests, or where tropical forests are degraded such that they open out to structurally resemble savannas, vegetation structure alone may be inadequate to distinguish mesic savanna from forest. Additional knowledge of the functional differences between these ecosystems which contrast sharply in their evolutionary and ecological history is required. Specifically, we suggest that tropical mesic savannas are predominantly mixed tree–C₄ grass systems defined by fire tolerance and shade intolerance of their species, while forests, from which C₄ grasses are largely absent, have species that are mostly fire intolerant and shade tolerant. Using this framework, we identify a suite of morphological, physiological and life‐history traits that are likely to differ between tropical mesic savanna and forest species. We suggest that these traits can be used to distinguish between these ecosystems and thereby aid their appropriate management and conservation. We also suggest that many areas in South Asia classified as tropical dry forests, but characterized by fire‐resistant tree species in a C₄ grass‐dominated understorey, would be better classified as mesic savannas requiring fire and light to maintain the unique mix of species that characterize them.     

Root dynamics influence tree–grass coexistence in an Australian savanna

Both resource and disturbance controls have been invoked to explain tree persistence among grasses in savannas. Here we determine the extent to which competition for available resources restricts the rooting depth of both grasses and trees, and how this may influence nutrient cycling under an infrequently burned savanna near Darwin, Australia. We sampled fine roots <2 mm in diameter from 24 soil pits under perennial as well as annual grasses and three levels of canopy cover. The relative proportion of C₃ (trees) and C₄ (grasses) derived carbon in a sample was determined using mass balance calculations. Our results show that regardless of the type of grass both tree and grass roots are concentrated in the top 20 cm of the soil. While trees have greater root production and contribute more fine root biomass grass roots contribute a disproportional amount of nitrogen and carbon to the soil relative to total root biomass. We postulate that grasses maintain soil nutrient pools and provide biomass for regular fires that prevent forest trees from establishing while savanna trees, are important for increasing soil N content, cycling and mineralization rates. We put forward our ideas as a hypothesis of resource‐regulated tree–grass coexistence in tropical savannas.     

Long-term effects of fire frequency and season on herbaceous vegetation in savannas of the Kruger National Park,South Africa

Aim s: The long-term effects of changing fire regimes on the herbaceous component of savannas are poorly understood but essential for understanding savanna dynamics. We present results from one of the longest running (>44 years) fire experiments in savannas, the experimental burn plots (EBPs), which is located in the Kruger National Park (South Africa) and encompasses four major savanna vegetation types that span broad spatial gradients of rainfall (450–700 mm) and soil fertility.Methods: Herbaceous vegetation was sampled twice in the EBPs using a modified step-point method, once prior to initiation of the experiment (1954) and again after 44–47 years. Different combinations of three fire frequency (1-, 2- and 3-year return intervals) and five season (before the first spring rains, after the first spring rains, mid-summer, late summer and autumn) treatments, as well as a fire exclusion treatment, were applied at the plot level (~7 ha each), with each treatment (n = 12 total) replicated four times at each of the four sites (n = 192 plots total). The effects of long-term alterations to the fire regime on grass community structure and composition were analyzed separately for each site.Important Findings: Over the 44+ years duration of the experiment, fires were consistently more intense on sites with higher mean annual rainfall (>570 mm), whereas fires were not as intense or consistent for sites with lower and more variable rainfall (<510 mm) and potentially higher herbivory due to greater soil fertility. Because the plots were open to grazing, the impacts of herbivory along with more variable rainfall regimes likely minimized the effects of fire for the more arid sites. As a consequence, fire effects on grass community structure and composition were most marked for the higher rainfall sites and generally not significant for the more arid sites. For the high-rainfall sites, frequent dry season fires (1- to 3-year return intervals) resulted in high grass richness, evenness and diversity, whereas fire exclusion and growing season fires had the lowest of these measures and diverged the most in composition as the result of increased abundance of a few key grasses. Overall, the long-term cumulative impacts of altered fire regimes varied across broad climatic and fertility gradients, with fire effects on the grass community decreasing in importance and herbivory and climatic variability likely having a greater influence on community structure and composition with increasing aridity and soil fertility.     

Effect of woody vegetation clearing on nutrient and carbon relations of semi-arid dystrophic savanna

Throughout the savanna biome, woody vegetation is cleared to increase productivity of herbaceous pasture. While clearing can result in increased pasture production of semi-arid dystrophic savannas in the short term, it is uncertain whether production is sustained in the long term. There is insufficient knowledge of how clearing affects soil nutrient and organic carbon (SOC) stocks. Using cleared-uncleared site pairs, we evaluated techniques for time-integrated assessment of nutrient and carbon relations in Australian savanna. Short-term in situ resin incubation showed that soil at cleared sites had a higher time-integrated availability of ammonium and nitrate, indicating that nitrogen (N) may turn over faster and/or is taken up slower at cleared sites than uncleared savanna. Nitrate and ammonium availability was approximately 2-fold higher in spring than in summer, likely due to greater uptake and/or loss of nitrate during summer rains. Nitrate was a prominent N source for evergreen trees, especially before summer rain, pointing to a role of trees as permanent N sinks. Stable isotope signatures of soil and vegetation indicate that N input occurs via N₂ fixing microbiotic crusts and Acacia species. 30 years after clearing, SOC contained more C₄ grass-derived carbon than uncleared savanna, but this shift in C source was not associated with the net C gain often observed in grasslands. Interactions between altered nutrient and C relations and composition of the understorey should be assessed in context of introduced buffelgrass (Cenchrus ciliaris) which had higher macronutrient concentrations than native grasses. Heterogeneity of the studied soils highlights the need for replication at several spatial scales to infer long-term dynamics with space-for-time chronosequences. We conclude that the techniques presented here are useful for gaining knowledge of the biogeochemical processes governing savannas and the systems that result from clearing.     

Long‐term variability and rainfall control of savanna fire regimes in equatorial East Africa

Fires burning the vast grasslands and savannas of Africa significantly influence the global carbon cycle. Projecting the impacts of future climate change on fire‐mediated biogeochemical processes in these dry tropical ecosystems requires understanding of how various climate factors influence regional fire regimes. To examine climate–vegetation–fire linkages in dry savanna, we conducted macroscopic and microscopic charcoal analysis on the sediments of the past 25 000 years from Lake Challa, a deep crater lake in equatorial East Africa. The charcoal‐inferred shifts in local and regional fire regimes were compared with previously published reconstructions of temperature, rainfall, seasonal drought severity, and vegetation dynamics to evaluate millennial‐scale drivers of fire occurrence. Our charcoal data indicate that fire in the dry lowland savanna of southeastern Kenya was not fuel‐limited during the Last Glacial Maximum (LGM) and Late Glacial, in contrast to many other regions throughout the world. Fire activity remained high at Lake Challa probably because the relatively high mean‐annual temperature (~22 °C) allowed productive C₄ grasses with high water‐use efficiency to dominate the landscape. From the LGM through the middle Holocene, the relative importance of savanna burning in the region varied primarily in response to changes in rainfall and dry‐season length, which were controlled by orbital insolation forcing of tropical monsoon dynamics. The fuel limitation that characterizes the region's fire regime today appears to have begun around 5000–6000 years ago, when warmer interglacial conditions coincided with prolonged seasonal drought. Thus, insolation‐driven variation in the amount and seasonality of rainfall during the past 25 000 years altered the immediate controls on fire occurrence in the grass‐dominated savannas of eastern equatorial Africa. These results show that climatic impacts on dry‐savanna burning are heterogeneous through time, with important implications for efforts to anticipate future shifts in fire‐mediated ecosystem processes.     

Contrasting demographics of tropical savanna and temperate forest eucalypts provide insight into how savannas and forests function. A case study using Corymbia clarksoniana from north‐eastern Australia

Eucalypts (Eucalyptus spp. and Corymbia spp.) dominate many communities across Australia, including frequently burnt tropical savannas and temperate forests, which receive less frequent but more intense fires. Understanding the demographic characteristics that allow related trees to persist in tropical savannas and temperate forest ecosystems can provide insight into how savannas and forests function, including grass–tree coexistence. This study reviews differences in critical stages in the life cycle of savanna and temperate forest eucalypts, especially in relation to fire. It adds to the limited data on tropical eucalypts, by evaluating the effect of fire regimes on the population biology of Corymbia clarksoniana, a tree that dominates some tropical savannas of north‐eastern Australia. Corymbia clarksoniana displays similar demographic characteristics to other tropical savanna species, except that seedling emergence is enhanced when seed falls onto recently burnt ground during a high rainfall period. In contrast to many temperate forest eucalypts, tropical savanna eucalypts lack canopy‐stored seed banks; time annual seed fall to coincide with the onset of predictable wet season rain; have very rare seedling emergence events, including a lack of mass germination after each fire; possess an abundant sapling bank; and every tropical eucalypt species has the ability to maintain canopy structure by epicormically resprouting after all but the most intense fires. The combination of poor seedling recruitment strategies, coupled with characteristics allowing long‐term persistence of established plants, indicate tropical savanna eucalypts function through the persistence niche rather than the regeneration niche. The high rainfall‐promoted seedling emergence of C. clarksoniana and the reduction of seedling survival and sapling growth by fire, support the predictions that grass–tree coexistence in savannas is governed by rainfall limiting tree seedling recruitment and regular fires limiting the growth of juvenile trees to the canopy.     

Herbaceous vegetation responses to experimental fire in savannas and forests depend on biome and climate

Fire–vegetation feedbacks potentially maintain global savanna and forest distributions. Accordingly, vegetation in savanna and forest ecosystems should have differential responses to fire, but fire response data for herbaceous vegetation have yet to be synthesized across biomes. Here, we examined herbaceous vegetation responses to experimental fire at 30 sites spanning four continents. Across a variety of metrics, herbaceous vegetation increased in abundance where fire was applied, with larger responses to fire in wetter and in cooler and/or less seasonal systems. Compared to forests, savannas were associated with a 4.8 (±0.4) times larger difference in herbaceous vegetation abundance for burned versus unburned plots. In particular, grass cover decreased with fire exclusion in savannas, largely via decreases in C₄ grass cover, whereas changes in fire frequency had a relatively weak effect on grass cover in forests. These differential responses underscore the importance of fire for maintaining the vegetation structure of savannas and forests.     

Woody cover in African savannas: the role of resources, fire and herbivory

Aim To determine the functional relationships between, and the relative importance of, different driver variables (mean annual precipitation, soil properties, fire and herbivory) in regulating woody plant cover across broad environmental gradients in African savannas. Location Savanna grasslands of East, West and Southern Africa. Methods The dependence of woody cover on mean annual precipitation (MAP), soil properties (texture, nitrogen mineralization potential and total phosphorus), fire regimes, and herbivory (grazer, browser + mixed feeder, and elephant biomass) was determined for 161 savanna sites across Africa using stochastic gradient boosting, a refinement of the regression tree analysis technique. Results All variables were significant predictors of woody cover, collectively explaining 71% of the variance in our data set. However, their relative importance as regulators of woody cover varied. MAP was the most important predictor, followed by fire return periods, soil characteristics and herbivory regimes. Woody cover showed a strong positive dependence on MAP between 200 and 700 mm, but no dependence on MAP above this threshold when the effects of other predictors were accounted for. Fires served to reduce woody cover below rainfall‐determined levels. Woody cover showed a complex, non‐linear relationship with total soil phosphorus, and was negatively correlated with clay content. There was a strong negative dependence of woody cover on soil nitrogen (N) availability, suggesting that increased N‐deposition may cause shifts in savannas towards more grassy states. Elephants, mixed feeders and browsers had negative effects on woody cover. Grazers, on the other hand, depressed woody cover at low biomass, but favoured woody vegetation when their biomass exceeded a certain threshold. Main conclusions Our results indicate complex and contrasting relationships between woody cover, rainfall, soil properties and disturbance regimes in savannas, and suggest that future environmental changes such as altered precipitation regimes, N‐enrichment and elevated levels of CO₂ are likely to have opposing, and potentially interacting, influences on the tree–grass balance in savannas.     

Biomass consumption by surface fires across Earth's most fire prone continent

Landscape fire is a key but poorly understood component of the global carbon cycle. Predicting biomass consumption by fire at large spatial scales is essential to understanding carbon dynamics and hence how fire management can reduce greenhouse gas emissions and increase ecosystem carbon storage. An Australia‐wide field‐based survey (at 113 locations) across large‐scale macroecological gradients (climate, productivity and fire regimes) enabled estimation of how biomass combustion by surface fire directly affects continental‐scale carbon budgets. In terms of biomass consumption, we found clear trade‐offs between the frequency and severity of surface fires. In temperate southern Australia, characterised by less frequent and more severe fires, biomass consumed per fire was typically very high. In contrast, surface fires in the tropical savannas of northern Australia were very frequent but less severe, with much lower consumption of biomass per fire (about a quarter of that in the far south). When biomass consumption was expressed on an annual basis, biomass consumed was far greater in the tropical savannas (>20 times that of the far south). This trade‐off is also apparent in the ratio of annual carbon consumption to net primary production (NPP). Across Australia's naturally vegetated land area, annual carbon consumption by surface fire is equivalent to about 11% of NPP, with a sharp contrast between temperate southern Australia (6%) and tropical northern Australia (46%). Our results emphasise that fire management to reduce greenhouse gas emissions should focus on fire prone tropical savanna landscapes, where the vast bulk of biomass consumption occurs globally. In these landscapes, grass biomass is a key driver of frequency, intensity and combustion completeness of surface fires, and management actions that increase grass biomass are likely to lead to increases in greenhouse gas emissions from savanna fires.     

Trees improve grass quality for herbivores in African savannas

The tree–grass interactions of African savannas are mainly determined by varying rainfall patterns and soil fertility. Large savanna trees are known to modify soil nutrient conditions, but whether this has an impact on the quality of herbaceous vegetation is unclear. However, if this were the case, then the removal of trees might also affect the structure and quality of the grass layer. We studied the impact of large nitrogen- and non-nitrogen fixing trees on the sub-canopy (SC) grass layer in low- and high-rainfall areas of differing soil fertility in eastern and southern Africa. We compared the structure and nutrient levels of SC grasses with those outside the canopy. Grass leaf nitrogen and phosphorus contents beneath tree canopies were elevated at all study sites and were up to 25% higher than those outside the canopy in the site of lowest rainfall and soil fertility. Grass leaf fibre and organic matter (OM) contents were slightly enhanced beneath tree canopies. At the site of highest rainfall and soil fertility, grasses beneath the canopy had significantly lower ratios of stem:leaf biomass and dead:living leaf material. Grass species composition differed significantly, with the highly nutritious Panicum spp. being most abundant underneath tree crowns. In the two drier study sites, soil nitrogen and OM contents were enhanced by 30% beneath trees. N-fixation capacity of trees did not contribute to the improved quality of grass under the canopy. We conclude that trees improve grass quality, especially in dry savannas. In otherwise nutrient-poor savanna grasslands, the greater abundance of high-quality grass species with higher contents of N and P and favourable grass structure beneath trees could attract grazing ungulates. As these benefits may be lost with tree clearance, trees should be protected in low fertility savannas and their benefits for grazing wildlife recognised in conservation strategies.     

Increased tree densities in South African savannas: >50 years of data suggests CO2 as a driver

For the past century, woody plants have increased in grasslands and savannas worldwide. Woody encroachment may significantly alter ecosystem functioning including fire regimes, herbivore carrying capacity, biodiversity and carbon storage capacity. Traditionally, increases in woody cover and density have been ascribed to changes in the disturbance regime (fire and herbivores) or rainfall. Increased atmospheric CO₂ concentrations may also contribute, by increasing growth rates of trees relative to grasses. This hypothesis is still heavily debated because usually potential CO₂ effects are confounded by changes in land use (disturbance regime). Here we analyse changes in woody density in fire experiments at three sites in South African savannas where the disturbance regime (fire and herbivores) was kept constant for 30 and 50 years. If global drivers had significant effects on woody plants, we would expect significant increases in tree densities and biomass over time under the constant disturbance regime. Woody density remained constant in a semiarid savanna but tripled in a mesic savanna between the 1970s and 1990s. At the third site, a semiarid savanna near the southern limits of the biome, tree density doubled from the mid 1990s to 2010. Interpretation of the causes is confounded by population recovery after clearing, but aerial photograph analysis on adjacent non‐cleared areas showed an accompanying 48% increase in woody cover. Increased CO₂ concentrations are consistent with increased woody density while other global drivers (rainfall) remained constant over the duration of the experiments. The absence of a response in one semiarid savanna could be explained by a smaller carbon sink capacity of the dominant species, which would therefore benefit less from increased CO₂. Understanding how savannas and grasslands respond to increased CO₂ and identifying the causes of woody encroachment are essential for the successful management of these systems.     

A patch-dynamics approach to savanna dynamics and woody plant encroachment – Insights from an arid savanna

The coexistence of woody and grassy plants in savannas has often been attributed to a rooting-niche separation (two-layer hypothesis). Water was assumed to be the limiting resource for both growth forms and grasses were assumed to extract water from the upper soil layer and trees and bushes from the lower layers. Woody plant encroachment (i.e. an increase in density of woody plants often unpalatable to domestic livestock) is a serious problem in many savannas and is believed to be the result of overgrazing in ‘two-layer systems’. Recent research has questioned the universality of both the two-layer hypothesis and the hypothesis that overgrazing is the cause of woody plant encroachment.

We present an alternative hypothesis explaining both tree–grass coexistence and woody plant encroachment in arid savannas. We propose that woody plant encroachment is part of a cyclical succession between open savanna and woody dominance and is driven by two factors: rainfall that is highly variable in space and time, and inter-tree competition. In this case, savanna landscapes are composed of many patches (a few hectares in size) in different states of transition between grassy and woody dominance, i.e. we hypothesize that arid savannas are patch-dynamic systems. We summarize patterns of tree distribution observed in an arid savanna in Namibia and show that these patterns are in agreement with the patch-dynamic savanna hypothesis. We discuss the applicability of this hypothesis to fire-dominated savannas, in which rainfall variability is low and fire drives spatial heterogeneity.

We conclude that field studies are more likely to contribute to a general understanding of tree–grass coexistence and woody plant encroachment if they consider both primary (rain and nutrients) and secondary (fire and grazing) determinants of patch properties across different savannas.     

Plant functional group responses to fire frequency and tree canopy cover gradients in oak savannas and woodlands

Questions: How do fire frequency, tree canopy cover, and their interactions influence cover of grasses, forbs and understorey woody plants in oak savannas and woodlands? Location: Minnesota, USA. Methods: We measured plant functional group cover and tree canopy cover on permanent plots within a long‐term prescribed fire frequency experiment and used hierarchical linear modeling to assess plant functional group responses to fire frequency and tree canopy cover. Results: Understorey woody plant cover was highest in unburned woodlands and was negatively correlated with fire frequency. C4‐grass cover was positively correlated with fire frequency and negatively correlated with tree canopy cover. C3‐grass cover was highest at 40% tree canopy cover on unburned sites and at 60% tree canopy cover on frequently burned sites. Total forb cover was maximized at fire frequencies of 4–7 fires per decade, but was not significantly influenced by tree canopy cover. Cover of N‐fixing forbs was highest in shaded areas, particularly on frequently burned sites, while combined cover of all other forbs was negatively correlated with tree canopy cover. Conclusions: The relative influences of fire frequency and tree canopy cover on understorey plant functional group cover vary among plant functional groups, but both play a significant role in structuring savanna and woodland understorey vegetation. When restoring degraded savannas, direct manipulation of overstorey tree canopy cover should be considered to rapidly reduce shading from fire‐resistant overstorey trees. Prescribed fires can then be used to suppress understorey woody plants and promote establishment of light‐demanding grasses and forbs.     

Variation in soil carbon stocks and their determinants across a precipitation gradient in West Africa

We examine the influence of climate, soil properties and vegetation characteristics on soil organic carbon (SOC) along a transect of West African ecosystems sampled across a precipitation gradient on contrasting soil types stretching from Ghana (15°N) to Mali (7°N). Our findings derive from a total of 1108 soil cores sampled over 14 permanent plots. The observed pattern in SOC stocks reflects the very different climatic conditions and contrasting soil properties existing along the latitudinal transect. The combined effects of these factors strongly influence vegetation structure. SOC stocks in the first 2 m of soil ranged from 20 Mg C ha^?1 for a Sahelian savanna in Mali to over 120 Mg C ha^?1 for a transitional forest in Ghana. The degree of interdependence between soil bulk density (SBD) and soil properties is highlighted by the strong negative relationships observed between SBD and SOC (r² > 0.84). A simple predictive function capable of encompassing the effect of climate, soil properties and vegetation type on SOC stocks showed that available water and sand content taken together could explain 0.84 and 0.86 of the total variability in SOC stocks observed to 0.3 and 1.0 m depth respectively. Used in combination with a suitable climatic parameter, sand content is a good predictor of SOC stored in highly weathered dry tropical ecosystems with arguably less confounding effects than provided by clay content. There was an increased contribution of resistant SOC to the total SOC pool for lower rainfall soils, this likely being the result of more frequent fire events in the grassier savannas of the more arid regions. This work provides new insights into the mechanisms determining the distribution of carbon storage in tropical soils and should contribute significantly to the development of robust predictive models of biogeochemical cycling and vegetation dynamics in tropical regions.     

Holocene boundary dynamics of a northern Australian monsoon rainforest patch inferred from isotopic analysis of carbon, (14C and δ13C) and nitrogen (δ15N) in soil organic matter

Abstract Soil organic matter (SOM) was sampled from lateritic soil profiles across an abrupt eucalypt savanna–monsoon rainforest boundary on the north coast of Croker Island, northern Australia. Accelerator mass spectrometry dating revealed that SOM that had accumulated at the base of these 1.5 m profiles had a radiocarbon age of about 5000 years. The mean carbon and nitrogen stable isotope composition of SOM from 10 cm deep layers from the surface, middle and base of three monsoon rainforest soil profiles was significantly different from the means for these layers in three adjacent savanna soil profiles, suggesting the isotopic ‘footprint’ of the vegetation boundary has been stable since the mid Holocene. Although there were no obvious environmental discontinuities associated with the boundary, the monsoon rainforest was found to occur on significantly more clay rich soils than the surrounding savanna. Tiny fragments of monsoon rainforest and abandoned ‘nests’ (large earthen mounds) of the orange‐footed scrubfowl, an obligate monsoon rainforest species, occurred in the savanna, signalling that the rainforest was once more extensive. Despite episodic disturbances, such as tropical storm damage and fires, the stability of the boundary is probably maintained because clay rich soils enable monsoon rainforest tree species to grow rapidly and achieve canopy closure, thereby excluding grass and reducing the risk of fire. Conversely, slower tree growth rates, grass competition and fire on the savanna soils would impede the expansion of the rainforest although high rainfall periods with shorter dry seasons may enable rainforest trees to grow sufficiently quickly to colonize the savanna successfully.     

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.     

Testing the grass-fire cycle: alien grass invasion in the tropical savannas of northern Australia

Abstract. Invasive alien grasses can increase fuel loads, leading to changes in fire regimes of invaded ecosystems by increasing the frequency, intensity and spatial extent of fires. Andropogon gayanus Kunth. (Gamba grass), a tall perennial grass from Africa, is invading ecosystems in the Top End of northern Australia. To determine whether A. gayanus alters savanna fire regimes, we compared fuel loads and fire intensities at invaded sites with those from native grass savannas. Savanna invaded by A. gayanus had fuel loads up to seven times higher than those dominated by native grasses. This higher fuel load supported a fire that was on average eight times more intense than those recorded in native grass savannas at the same time of year (means 15700 ± 6200 and 2100 ± 290 kW m⁻¹, respectively), and was the highest early dry season fire intensities ever recorded in the Northern Territory. These results suggest that A. gayanus is a serious threat to northern Australia's savannas, with the potential to alter vegetation structure and initiate a grass-fire cycle.     

Optimal Fire Regimes for Soil Carbon Storage in Tropical Savannas of Northern Australia

Modification of fire regimes in tropical savannas can have significant impacts on the global carbon (C) cycle, and therefore, on the climate system. In Australian tropical savannas, there has been recent, large-scale implementation of fire management that aims to decrease Kyoto-compliant non-CO₂ greenhouse gas emissions by reducing late dry season intense fires through strategic early dry season burning. However, there is no accounting for changes to soil C stocks resulting from changes to savanna fire management, although impacts on these pools may be considerable. We present a hypothesis that soil C storage is greatest under low intensity fires with an intermediate fire return interval. Simulations using the CENTURY Soil Organic Matter Model confirmed this hypothesis with greatest soil C storage under a fire regime of one low intensity fire every 5 years. Key areas of uncertainty for CENTURY model simulations include fine root dynamics, charcoal production and nitrogen (N) cycling, and better understanding of these processes could improve model predictions. Soil C stocks measured in the field after 5 years of annual, 3 year and unburned fire treatments were not significantly different (range 41–58 t ha⁻¹), but further CENTURY modelling suggests that changes in fire management will take up to 100 years to have a detectable impact (+4 t ha⁻¹) on soil C stocks. However, implementation of fire management that reduces fire frequency and intensity within the large area of intact savanna landscapes in northern Australia could result in emissions savings of 0.17 t CO₂-e ha⁻¹ y⁻¹, four times greater than reductions of non-CO₂ emissions.     

Mammalian herbivores,grass height and rainfall drive termite activity at different spatial scales in an African savanna

Termites have a large influence on ecosystem functioning. Understanding what drives termite activity patterns improves understanding of nutrient cycling, productivity, and heterogeneity in savannas. We present a mechanistic framework that relates the interactive effects of rainfall, grassland structure, large herbivore presence, and soil factors to termite activity. To test this framework, we used grass litterbags to monitor termite activity at ten sites across Hluhluwe‐iMfolozi Park, South Africa. We assessed the effects of abiotic and biotic factors on termite activity at two scales: the large (landscape) scale, variation in bait removal among 300 m² plots that were distributed across the park and at the small (within‐plot) scale (1–300 m²). Half of our sites were located inside large herbivore exclosures to test for the effect of mammalian herbivore presence. At the landscape scale, termite grass removal declined towards higher rainfall and in the presence of mammalian herbivores. Removal did not depend on soil factors. At the small scale, removal declined with increasing grass height, particularly in the 1 m surrounding the bait bag. Resource quality did not affect bait removal. We suggest that competition for forage drives the negative effect of mammalian herbivores on termites, whereas lower bait removal in taller swards may be due to direct negative effects from rainfall, fire and/or competition with free‐living microbes. Ultimately, we suggest that the impact of termites on nutrient cycling is most pronounced when abiotic (rainfall) and biotic conditions (mammalian herbivory) limit grass removal by fire and decomposition by free‐living microbes.     

Decadal dynamics of tree cover in an Australian tropical savanna

Spatio‐temporal variation in tropical savanna tree cover remains poorly understood. We aimed to quantify the drivers of tree cover in tropical mesic savannas in Kakadu National Park by relating changes in tree cover over 40 years to: mean annual rainfall, fire activity, initial tree cover and prior changes in tree cover. Aerial photography, acquired in 1964, 1984 and 2004, was obtained for fifty sites in Kakadu that spanned a rainfall gradient from approximately 1200 to 1600 mm. The remotely sensed estimates of tree cover were validated via field survey. Linear mixed effects modelling and multi‐model inference were used to assess the strength and form of the relationships between tree cover and predictor variables. Over the 40 years, tree cover across these savannas increased on average by 4.94 ± 0.88%, but was spatio‐temporally variable. Tree cover showed a positive albeit weak trend across the rainfall gradient. The strength of this positive relationship varied over the three measurement times, and this suggests that other factors are important in controlling tree cover. Tree cover was positively related to prior tree cover, and negatively correlated with fire activity. Over 20 years tree cover was more likely to increase if (i) tree cover was initially low or (ii) had decreased in the previous 20‐year interval or (iii) there had been fewer fires. Across the examined rainfall gradient, the greater variability in fire activity and inherently higher average tree cover at the wetter latitudes resulted in greater dynamism of tree cover compared with the drier latitudes. This is consistent with savanna tree cover being determined by interactions between mean annual rainfall, tree competition and frequent fire in these tropical mesic savannas.