Navashni Govender

EFFECTS OF FOUR DECADES OF FIRE MANIPULATION ON WOODY VEGETATION STRUCTURE IN SAVANNA

The amount of carbon stored in savannas represents a significant uncertainty in global carbon budgets, primarily because fire causes actual biomass to differ from potential biomass. We analyzed the structural response of woody plants to long-term experimental burning in savannas. The experiment uses a randomized block design to examine fire exclusion and the season and frequency of burn in 192 7-ha experimental plots located in four different savanna ecosystems. Although previous studies would lead us to expect tree density to respond to the fire regime, our results, obtained from four different savanna ecosystems, suggest that the density of woody individuals was unresponsive to fire. The relative dominance of small trees was, however, highly responsive to fire regime. The observed shift in the structure of tree populations has potentially large impacts on the carbon balance. However, the response of tree biomass to fire of the different savannas studied were different, making it difficult to generalize about the extent to which fire can be used to manipulate carbon sequestration in savannas. This study provides evidence that savannas are demographically resilient to fire, but structurally responsive.

Effects of fire on woody vegetation structure in African savanna.

Despite the importance of fire in shaping savannas, it remains poorly understood how the frequency, seasonality, and intensity of fire interact to influence woody vegetation structure, which is a key determinant of savanna biodiversity. We provide a comprehensive analysis of vertical and horizontal woody vegetation structure across one of the oldest savanna fire experiments, using new airborne Light Detection and Ranging (LiDAR) technology. We developed and compared high-resolution woody vegetation height surfaces for a series of large experimental burn plots in the Kruger National Park, South Africa. These 7-ha plots (total area approximately 1500 ha) have been subjected to fire in different seasons and at different frequencies, as well as no-burn areas, for 54 years. Long-term exposure to fire caused a reduction in woody vegetation up to the 5.0-7.5 m height class, although most reduction was observed up to 4 m. Average fire intensity was positively correlated with changes in woody vegetation structure. More frequent fires reduced woody vegetation cover more than less frequent fires, and dry-season fires reduced woody vegetation more than wet-season fires. Spring fires from the late dry season reduced woody vegetation cover the most, and summer fires from the wet season reduced it the least. Fire had a large effect on structure in the densely wooded granitic landscapes as compared to the more open basaltic landscapes, although proportionally, the woody vegetation was more reduced in the drier than in the wetter landscapes. We show that fire frequency and fire season influence patterns of vegetation three-dimensional structure, which may have cascading consequences for biodiversity. Managers of savannas can therefore use fire frequency and season in concert to achieve specific vegetation structural objectives.

The contribution of fire research to fire management: a critical review of a long-term experiment in the Kruger National Park, South Africa

The present paper reviews a long-term fire experiment in the Kruger National Park, South Africa, established in 1954 to support fire management. The paper’s goals are: (1) to assess learning, with a focus on relevance for fire management; (2) to examine how findings influenced changes in fire management; and (3) to reflect on the experiment’s future. Results show that fire treatments affected vegetation structure and biomass more than species composition. Effects on vegetation were most marked in extreme treatments (annual burning, burning in the summer wet season, or long periods of fire exclusion), and were greater in areas of higher rainfall. Faunal communities and soil physiology were largely unaffected by fire. Since the inception of the experiment, paradigms in savanna ecology have changed to encompass heterogeneity and variability. The design of the experiment, reflecting the understanding of the 1950s, does not cater for variability, and as a result, the experiment had little direct influence on changes in management policy. Notwithstanding this, managers accept that basic research influences the understanding of fundamental ecosystem function, and they recognise that it promotes appropriate adaptive management by contributing to predictive understanding. This has been a major reason for maintaining the experiment for over 50 years.

Controls of aboveground net primary production in mesic savanna grasslands: an inter-hemispheric comparison.

Patterns and controls of annual aboveground net primary productivity (ANPP) are fundamental metrics of ecosystem functioning. It is generally assumed, but rarely tested, that determinants of ANPP in one region within a biome will operate similarly throughout that biome, as long as physiognomy and climate are broadly consistent. We tested this assumption by quantifying ANPP responses to fire, grazing history, and nitrogen (N) addition in North American (NA) and South African (SA) savanna grasslands. We found that total ANPP responded in generally consistent ways to fire, grazing history, and N addition on both continents. Annual fire in both NA and SA consistently stimulated total ANPP (28–100%) relative to unburned treatments at sites with deep soils, and had no effect on ANPP in sites with shallow soils. Fire did not affect total ANPP in sites with a recent history of grazing, regardless of whether a single or a diverse suite of large herbivores was present. N addition interacted strongly and consistently with fire regime in both NA and SA. In annually burned sites that were not grazed, total ANPP was stimulated by N addition (29–39%), but there was no effect of N fertilization in the absence of fire. In contrast, responses in forb ANPP to fire and grazing were somewhat divergent across this biome. Annual fire in NA reduced forb ANPP, whereas grazing increased forb ANPP, but neither response was evident in SA. Thus, despite a consistent response in total ANPP, divergent responses in forb ANPP suggest that other aspects of community structure and ecosystem functioning differ in important ways between these mesic savanna grasslands.

Fire alters ecosystem carbon and nutrients but not plant nutrient stoichiometry or composition in tropical savanna

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 ...

Plant community response to loss of large herbivores differs between North American and South African savanna grasslands

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.

Loss of a large grazer impacts savanna grassland plant communities similarly in North America and South Africa

Abstract Large herbivore grazing is a widespread disturbance in mesic savanna grasslands which increases herbaceous plant community richness and diversity. However, humans are modifying the impacts of grazing on these ecosystems by removing grazers. A more general understanding of how grazer loss will impact these ecosystems is hampered by differences in the diversity of large herbivore assemblages among savanna grasslands, which can affect the way that grazing influences plant communities. To avoid this we used two unique enclosures each containing a single, functionally similar large herbivore species. Specifically, we studied a bison (Bos bison) enclosure at Konza Prairie Biological Station, USA and an African buffalo (Syncerus caffer) enclosure in Kruger National Park, South Africa. Within these enclosures we erected exclosures in annually burned and unburned sites to determine how grazer loss would impact herbaceous plant communities, while controlling for potential fire-grazing interactions. At both sites, removal of the only grazer decreased grass and forb richness, evenness and diversity, over time. However, in Kruger these changes only occurred with burning. At both sites, changes in plant communities were driven by increased dominance with herbivore exclusion. At Konza, this was caused by increased abundance of one grass species, Andropogon gerardii, while at Kruger, three grasses, Themeda triandra, Panicum coloratum, and Digitaria eriantha increased in abundance.

Woody plant biomass and carbon exchange depend on elephant‐fire interactions across a productivity gradient in African savanna

Elephants and fire are individually well-known disturbance agents within savanna ecosystems, but their interactive role in governing tree-cover dynamics and savanna-forest biome boundaries remain unresolved. Of central importance are the mechanisms by which elephants vs. fire affect tree biomass and cover, and how – over long time periods – both factors interact with rainfall and soils to govern tree biomass and carbon dynamics. Here, we evaluated the response of woody vegetation to 56 years of fire manipulation in South Africas Kruger National Park, with three fire regimes (annual, triennial, and unburned) replicated across a productivity gradient and subject to two periods of contrasting elephant abundances (generated by the cessation of culling in 1994). Higher fire frequencies had a negative effect on woody biomass in the low-elephant period, but this effect was weak-to-negligible in the high-elephant period as the difference among fire treatments diminished. Moreover, elephants removed increasing amounts of woody biomass as productivity increased across study sites, but fire did not. We infer that elephant-induced tree mortality could overcome increases in woody-plant productivity, while fire-induced mortality alone could not. Elephants caused woody-plant carbon to shift from a sink to a source; this effect was independent of fire treatment, with highest rates of net carbon removal in the wettest and most productive site. Synthesis: Our results reveal a context-dependent interaction between fire and elephants as disturbance agents in savanna: the influence of fire on woody plants was sensitive to the abundance of elephants and diminished with increased plant productivity. In contrast, elephants were capable of shifting landscapes from relatively dense woodland to open savanna, even in unburned sites, and exerted strong impacts irrespective of site conditions and plant productivity. This article is protected by copyright. All rights reserved.

Fire Alters Ecosystem Carbon and Nutrients but not Plant Nutrient Stoichiometry

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.