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Summary Frequent fires are often proposed as a way of preventing woody encroachment in savannas, yet few studies have examined whether high‐intensity fires can effectively reverse woody encroachment. We applied successive fire treatments to examine the effect of fire intensity on woody vegetation structure. The treatments included early dry season, low‐intensity fires; late dry season, higher‐intensity fires; and an unburnt control. We used pre‐ and post‐fire airborne LiDAR to compare vegetation structural changes brought about by fires of different intensity. Early dry season fires were of lower intensity (1400–2100 kW m−1) than late dry season fires (2500–4300 kW m−1). The two treatments also differed in terms of fuel consumed, scorch heights and char heights, indicating that clear differences in fire intensity and severity were achieved. After 4 years and two fire applications, relative woody cover increased by between 20 and 110% in different height categories following low‐intensity and control treatments and declined by between 3 and 70% following high‐intensity fire treatments. Declines were markedly higher following two repeated high‐intensity fires than following a high and then a moderate‐intensity fire. Because woody shrubs in lower height classes can recover rapidly, repeated high‐intensity fires would be needed to maintain lower cover. Tall trees are often assumed to be unaffected by fires. However, we found that the rate of tree loss was directly related to fire intensity, where 36% of trees were lost following repeated high‐intensity fires, compared to 22% after a high‐ and then a moderate‐intensity fire and 6% after two low‐intensity fires (3% without fire). Synthesis and applications. Using LiDAR data we show that high‐intensity fires can, at least in the short term, significantly reduce woody cover in South African savannas. The use of repeated high‐intensity fires simultaneously causes both a positive (reduction in cover of short shrubs) and a negative (loss of tall trees) outcome, and managers need to make trade‐offs when contemplating the use of fire intensity to achieve specific goals. One potential solution may be to repeatedly apply high‐intensity treatments to some areas, and not to others. This could generate a heterogeneous landscape where grasses become dominant and tall trees become scarce in some places, but in others, tall trees persist (or at least decline at slower rates), and shorter woody shrubs increase in dominance. Whether this would be acceptable, or practical, remains to be tested. Using LiDAR data we show that high‐intensity fires can, at least in the short term, significantly reduce woody cover in South African savannas. The use of repeated high‐intensity fires simultaneously causes both a positive (reduction in cover of short shrubs) and a negative (loss of tall trees) outcome, and managers need to make trade‐offs when contemplating the use of fire intensity to achieve specific goals. One potential solution may be to repeatedly apply high‐intensity treatments to some areas, and not to others. This could generate a heterogeneous landscape where grasses become dominant and tall trees become scarce in some places, but in others, tall trees persist (or at least decline at slower rates), and shorter woody shrubs increase in dominance. Whether this would be acceptable, or practical, remains to be tested.

Fire and nutrients interact to influence the global distribution and dynamics of the savanna biome, but the results of these interactions are both complex and poorly known. A critical but unresolved question is whether short-term losses of carbon and nutrients caused by fire can trigger long-term and potentially compensatory responses in the nutrient stoichiometry of plants, or in the abundance of dinitrogen-fixing trees. There is disagreement in the literature about the potential role of fire on savanna nutrients, and, in turn, on plant stoichiometry and composition. A major limitation has been the lack of fire manipulations over time scales sufficiently long for these interactions to emerge. We use a 58-year, replicated, large-scale, fire manipulation experiment in Kruger National Park (South Africa) in savanna to quantify the effect of fire on (1) distributions of carbon, nitrogen, and phosphorus at the ecosystem scale; (2) carbon : nitrogen : phosphorus stoichiometry of above- and belowground tissues of plant species; and (3) abundance of plant functional groups including nitrogen fixers. Our results show dramatic effects of fire on the relative distribution of nutrients in soils, but that individual plant stoichiometry and plant community composition remained unexpectedly resilient. Moreover, measures of nutrients and carbon stable isotopes allowed us to discount the role of tree cover change in favor of the turnover of herbaceous biomass as the primary mechanism that mediates a transition from low to high soil carbon and nutrients in the absence of fire. We conclude that, in contrast to extra-tropical grasslands or closed-canopy forests, vegetation in the savanna biome may be uniquely adapted to nutrient losses caused by recurring fire.

For large herbivores living in highly dynamic environments, maintaining range fidelity has the potential to facilitate the exploitation of predictable resources while minimising energy expenditure. We evaluate this expectation by examining how the seasonal range fidelity of African elephants ( Loxodonta africana ) in the Kruger National Park, South Africa is affected by spatiotemporal variation in environmental conditions (vegetation quality, temperature, rainfall, and fire). Eight-years of GPS collar data were used to analyse the similarity in seasonal utilisation distributions for thirteen family groups. Elephants exhibited remarkable consistency in their seasonal range fidelity across the study with rainfall emerging as a key driver of space-use. Within years, high range fidelity from summer to autumn and from autumn to winter was driven by increased rainfall and the retention of high-quality vegetation. Across years, sequential autumn seasons demonstrated the lowest levels of range fidelity due to inter-annual variability in the wet to dry season transition, resulting in unpredictable resource availability. Understanding seasonal space use is important for determining the effects of future variability in environmental conditions on elephant populations, particularly when it comes to management interventions. Indeed, over the coming decades climate change is predicted to drive greater variability in rainfall and elevated temperatures in African savanna ecosystems. The impacts of climate change also present particular challenges for elephants living in fragmented or human-transformed habitats where the opportunity for seasonal range shifts are greatly constrained.

Savannas cover 60% of the land surface in Southern Africa, with fires and herbivory playing a key role in their ecology. The Limpopo National Park (LNP) is a 10,000 km2 conservation area in southern Mozambique and key to protecting savannas in the region. Fire is an important factor in LNP's landscapes, but little is known about its role in the park's ecology. In this study, we explored the interaction between fire frequency (FF), landscape type, and vegetation. To assess the FF, we analyzed ten years of the Moderate resolution Imaging Spectroradiometer (MODIS) burned area product (2003–2013). A stratified random sampling approach was used to assess biodiversity across three dominant landscapes (Nwambia Sandveld‐NS, Lebombo North‐LN, and Shrubveld Mopane on Calcrete‐C) and two FF levels (low—twice or less; and high—3 times or more, during 10 years). Six ha were sampled in each stratum, except for the LN versus high FF in which low accessibility allowed only 3 ha sampling. FF was higher in NS and LN landscapes, where 25% and 34% of the area, respectively, burned more than three times in 10 years. The landscape type was the main determinant of grass composition and biomass. However, in the sandy NS biomass was higher under high FF. The three landscapes supported three different tree/shrub communities, but FF resulted in compositional variations in NS and LN. Fire frequency had no marked influence on woody structural parameters (height, density, and phytomass). We concluded that the savannas in LNP are mainly driven by landscape type (geology), but FF may impose specific modifications. We recommend a fire laissez‐faire management system for most of the park and a long‐term monitoring system of vegetation to address vegetation changes related to fire. Fire management should be coordinated with the neighboring Kruger National Park, given its long history of fire management. Synthesis: This study revealed that grass and tree/shrub density, biomass, and composition in LNP are determined by the landscape type, but FF determines some important modifications. We conclude that at the current levels FF is not dramatically affecting the savanna ecosystem in the LNP (Figure 1). However, an increase in FF may drive key ecosystem changes in grass biomass and tree/shrub species composition, height, phytomass, and density. This study revealed that grass and tree/shrub density, biomass, and composition in LNP are determined by the landscape type, but FF determines some important modifications. We conclude that at the current levels FF is not dramatically affecting the savanna ecosystem in the LNP (Figure 1). However, an increase in FF may drive key ecosystem changes in grass biomass and tree/shrub species composition, height, phytomass, and density.

Herbivory and fire shape plant community structure in grass-dominated ecosystems, but these disturbance regimes are being altered around the world. To assess the consequences of such alterations, we excluded large herbivores for seven years from mesic savanna grasslands sites burned at different frequencies in North America (Konza Prairie Biological Station, Kansas, USA) and South Africa (Kruger National Park). We hypothesized that the removal of a single grass-feeding herbivore from Konza would decrease plant community richness and shift community composition due to increased dominance by grasses. Similarly, we expected grass dominance to increase at Kruger when removing large herbivores, but because large herbivores are more diverse, targeting both grasses and forbs, at this study site, the changes due to herbivore removal would be muted. After seven years of large-herbivore exclusion, richness strongly decreased and community composition changed at Konza, whereas little change was evident at Kruger. We found that this divergence in response was largely due to differences in the traits and numbers of dominant grasses between the study sites rather than the predicted differences in herbivore assemblages. Thus, the diversity of large herbivores lost may be less important in determining plant community dynamics than the functional traits of the grasses that dominate mesic, disturbance-maintained savanna grasslands.

Herbivores are a main driver of ecosystem patterns and processes in semi-arid savannas, with their effects clearly observed when they are excluded from landscapes. Starting in the 1960s, various herbivore exclosures have been erected in the Kruger National Park (KNP), for research and management purposes. These exclosures vary from very small (1 m2) to relatively large (almost 900 ha), from short-term (single growing season) to long-term (e.g. some of the exclosures were erected more than 60 years ago), and are located on different geologies and across a rainfall gradient. We provide a summary of the history and specifications of various exclosures. This is followed by a systematic overview of mostly peer-reviewed literature resulting from using KNP exclosures as research sites. These 75 articles cover research on soils, vegetation dynamics, herbivore exclusion on other faunal groups and disease. We provide general patterns and mechanisms in a synthesis section, and end with recommendations to increase research outputs and productivity for future exclosure experiments.Conservation ImplicationsHerbivore exclosures in the KNP have become global research platforms, that have helped in the training of ecologists, veterinarians and field biologists, and have provided valuable insights into savanna dynamics that would otherwise have been hard to gain. In an age of dwindling conservation funding, we make the case for the value added by exclosures and make recommendations for their continued use as learning tools in complex African savannas.

To pursue the development and validation of coupled fire-atmosphere models, the wildland fire modeling community needs validation data sets with scenarios where fire-induced winds influence fire front behavior, and with high temporal and spatial resolution. Helicopter-borne infrared thermal cameras have the potential to monitor landscape-scale wildland fires at a high resolution during experimental burns. To extract valuable information from those observations, three-step image processing is required: (a) Orthorectification to warp raw images on a fixed coordinate system grid, (b) segmentation to delineate the fire front location out of the orthorectified images, and (c) computation of fire behavior metrics such as the rate of spread from the time-evolving fire front location. This work is dedicated to the first orthorectification step, and presents a series of algorithms that are designed to process handheld helicopter-borne thermal images collected during savannah experimental burns. The novelty in the approach lies on its recursive design, which does not require the presence of fixed ground control points, hence relaxing the constraint on field of view coverage and helping the acquisition of high-frequency observations. For four burns ranging from four to eight hectares, long-wave and mid infra red images were collected at 1 and 3 Hz, respectively, and orthorectified at a high spatial resolution (<1 m) with an absolute accuracy estimated to be lower than 4 m. Subsequent computation of fire radiative power is discussed with comparison to concurrent space-borne measurements.

In areas with diverse herbivore communities such as African savannas, the frequency of disturbance by fire may alter the top–down role of different herbivore species on plant community dynamics. In a seven year experiment in the Kruger National Park, South Africa, we examined the habitat use of nine common herbivore species across annually burned, triennially burned and unburned areas. We also used two types of exclosures (plus open access controls) to examine the impacts of different herbivores on plant community dynamics across fire disturbance regimes. Full exclosures excluded all herbivores > 0.5 kg (e.g. elephant, zebra, impala) while partial exclosures allowed access only to animals with shoulder heights ≤ 0.85 m (e.g. impala, steenbok). Annual burns attracted a diverse suite of herbivores, and exclusion of larger herbivores (e.g. elephant, zebra, wildebeest) increased plant abundance. When smaller species, mainly impala, were also excluded there were declines in plant diversity, likely mediated by a decline in open space available for colonization of uncommon plant species. Unburned areas attracted the least diverse suite of herbivores, dominated by impala. Here, herbivore exclusion, especially of impala, led to strong declines in plant richness and diversity. With no fire disturbance, herbivore exclusion led to competitive exclusion via increases in plant dominance and light limitation. In contrast, on triennial burns, herbivore exclusion had no effect on plant richness or diversity, potentially due to relatively little open space for colonization across exclosure treatments but also little competitive exclusion due to the intermediate fire disturbance. Further, the diverse suite of grazers and browsers on triennial burns may have had a compensating effect of on the diversity of grasses and forbs. Ultimately, our work shows that differential disturbance regimes can result in differential consumer pressure across a landscape and result in heterogeneous patterns in top–down control of community dynamics.

Fire is considered a critical management tool in fire prone landscapes. Often studies and policies relating to fire focus on why and how the fire regime should be managed, often neglecting to subsequently evaluate management's ability to achieve these objectives over long temporal and large spatial scales. This study explores to what extent the long-term spatio-temporal fire patterns recorded in the Kruger National Park, South Africa has been influenced by management policies and to what extent it was dictated by underlying variability in the abiotic template. This was done using a spatially explicit fire-scar database from 1941 to 2006 across the 2 million ha Park. Fire extent (ha burnt per annum) 1) is correlated with rainfall cycles 2) exhibits no long-term trend and 3) is largely non-responsive to prevailing fire management policies. Rainfall, geology and distance from the closest perennial river and the interactions between these variables influence large-scale fire pattern heterogeneity: areas with higher rainfall, on basaltic substrates and far from rivers are more fire prone and have less heterogeneous fire regimes than areas with lower rainfall, on granitic substrates and closer to rivers. This study is the first to illustrate that under a range of rainfall and geological conditions, perennial rivers influence longterm, landscape-scale fire patterns well beyond the riparian zone (typically up to 15 km from the river). It was concluded that despite fire management policies which historically aimed for largely homogeneous fire return regimes, spatially and temporally heterogeneous patterns have emerged. This is primarily because of differences in rainfall, geology and distance from perennial rivers. We postulate that large-scale spatio-temporal fire pattern heterogeneity is implicit to heterogeneous savannas, even under largely homogenizing fire policies. Management should be informed by these patterns, embracing the natural heterogeneity-producing template. We therefore suggest that management actions will be better directed when operating at appropriate scales, nested within the broader implicit landscape patterns, and when focusing on fire regime parameters over which they have more influence (e.g. fire season).

Alien plants were first recorded in 1937 in the 2 million ha Kruger National Park (KNP, a savanna protected area in South Africa), and attempts to control them began in the mid-1950s. The invasive alien plant control program expanded substantially in the late 1990s, but its overall efficacy has not been determined. We present an assessment of invasive alien plant control operations over several decades in KNP. We based our assessment on available information from a range of control programs funded from various sources, including national public works programs, KNP operational funds, and foreign donor funds. Over ZAR 350 million (~ US$ 27 million) has been spent on control interventions between 1997 and 2016. We found evidence of good progress with the control of several species, notably Opuntia stricta , Sesbania punicea , Lantana camara and several aquatic weeds, often because of effective biological control. On the other hand, we found that over one third (40%) of the funding was spent on species that have subsequently been recognised as being of lower priority, most of which were alien annual weeds. The allocation of funds to non-priority species was sometimes driven by the need to meet additional objectives (such as employment creation), or by perceptions about relative impact in the absence of documented evidence. We also found that management goals were limited to inputs (funds disbursed, employment created, and area treated) rather than to ecological outcomes, and progress was consequently not adequately monitored. At a species level, four out of 36 species were considered to be under complete control, and a further five were under substantial control. Attempts to control five annual species were all considered to be ineffective. On the basis of our findings, we recommend that more studies be done to determine impacts associated with individual invasive alien species; that the criteria used to prioritise invasive alien species be documented based on such assessments, so that management can justify a focus on priority species; and that funding be re-directed to those species that clearly pose greater threats, and for which other solutions (such as biological control) are not an option.