Will trees or grasses profit from changing rainfall regimes in savannas?

Increasing rainfall variability is widely expected under future climate change scenarios. How will savanna trees and grasses be affected by growing season dry spells and altered seasonality and how tightly coupled are tree-grass phenologies with rainfall? We measured tree and grass responses to growing season dry spells and dry season rainfall. We also tested whether the phenologies of 17 deciduous woody species and the Soil Adjusted Vegetation Index of grasses were related to rainfall between 2019 and 2023. Tree and grass growth was significantly reduced during growing season dry spells. Tree growth was strongly related to growing season soil water potentials and limited to the wet season. Grasses can rapidly recover after growing season dry spells and grass evapotranspiration was significantly related to soil water potentials in both the wet and dry seasons. Tree leaf flushing commenced before the rainfall onset date with little subsequent leaf flushing. Grasses grew when moisture became available regardless of season. Our findings suggest that increased dry spell length and frequency in the growing season may slow down tree growth in some savannas, which together with longer growing seasons may allow grasses an advantage over C3 plants that are advantaged by rising CO2 levels.

Quantifying the environmental limits to fire spread in grassy ecosystems.

Modeling fire spread as an infection process is intuitive: An ignition lights a patch of fuel, which infects its neighbor, and so on. Infection models produce nonlinear thresholds, whereby fire spreads only when fuel connectivity and infection probability are sufficiently high. These thresholds are fundamental both to managing fire and to theoretical models of fire spread, whereas applied fire models more often apply quasi-empirical approaches. Here, we resolve this tension by quantifying thresholds in fire spread locally, using field data from individual fires (n = 1,131) in grassy ecosystems across a precipitation gradient (496 to 1,442 mm mean annual precipitation) and evaluating how these scaled regionally (across 533 sites) and across time (1989 to 2012 and 2016 to 2018) using data from Kruger National Park in South Africa. An infection model captured observed patterns in individual fire spread better than competing models. The proportion of the landscape that burned was well described by measurements of grass biomass, fuel moisture, and vapor pressure deficit. Regionally, averaging across variability resulted in quasi-linear patterns. Altogether, results suggest that models aiming to capture fire responses to global change should incorporate nonlinear fire spread thresholds but that linear approximations may sufficiently capture medium-term trends under a stationary climate.

Limited increases in savanna carbon stocks over decades of fire suppression.

Savannas cover a fifth of the land surface and contribute a third of terrestrial net primary production, accounting for three-quarters of global area burned and more than half of global fire-driven carbon emissions1-3. Fire suppression and afforestation have been proposed as tools to increase carbon sequestration in these ecosystems2,4. A robust quantification of whole-ecosystem carbon storage in savannas is lacking however, especially under altered fire regimes. Here we provide one of the first direct estimates of whole-ecosystem carbon response to more than 60 years of fire exclusion in a mesic African savanna. We found that fire suppression increased whole-ecosystem carbon storage by only 35.4 ± 12% (mean ± standard error), even though tree cover increased by 78.9 ± 29.3%, corresponding to total gains of 23.0 ± 6.1 Mg C ha-1 at an average of about 0.35 ± 0.09 Mg C ha-1 year-1, more than an order of magnitude lower than previously assumed4. Frequently burned savannas had substantial belowground carbon, especially in biomass and deep soils. These belowground reservoirs are not fully considered in afforestation or fire-suppression schemes but may mean that the decadal sequestration potential of savannas is negligible, especially weighed against concomitant losses of biodiversity and function.

Woody encroachment happens via intensification, not extensification, of species ranges in an African savanna.

Widespread woody encroachment is a prominent concern for savanna systems as it is often accompanied by losses in productivity and biodiversity. Extensive ecosystem-level work has advanced our understanding of its causes and consequences. However, there is still debate over whether local management can override regional and global drivers of woody encroachment, and it remains largely unknown how encroachment influences woody community assemblages. Here, we examined species-level changes in woody plant distributions and size structure from the late 1980s to the late 2000s based on spatially intensive ground-based surveys across Kruger National Park, South Africa. This study region spans broad gradients in rainfall, soil texture, fire frequency, elephant density, and other topographic variables. Species-level changes in frequency of occurrence and size class proportion reflected widespread woody encroachment primarily by Dichrostachys cinerea and Combretum apiculatum, and a loss of large trees mostly of Sclerocarya birrea and Acacia nigrescens. Environmental variables determining woody species distributions across Kruger varied among species but did not change substantially between two sampling times, indicating that woody encroachers were thickening within their existing ranges. Overall, more areas across Kruger were found to have an increased number of common woody species through time, which indicated an increase in stem density. These areas were generally associated with decreasing fire frequency and rainfall but increasing elephant density. Our results suggest that woody encroachment is a widespread but highly variable trend across landscapes in Kruger National Park and potentially reflects an erosion of local heterogeneity in woody community assemblages. Many savanna managers, including in Kruger, aim to manage for heterogeneity in order to promote biodiversity, where homogenization of vegetation structure counters this specific goal. Increasing fire frequency has some potential as a local intervention. However, many common species increased in commonness even under near-constant disturbance conditions, which likely limits the potential for managing woody encroachment in the face of drivers beyond the scope of local control. Regular field sampling coupled with targeted fire management will enable more accurate monitoring of the rate of encroachment intensification.

Decadal changes in fire frequencies shift tree communities and functional traits.

Global change has resulted in chronic shifts in fire regimes. Variability in the sensitivity of tree communities to multi-decadal changes in fire regimes is critical to anticipating shifts in ecosystem structure and function, yet remains poorly understood. Here, we address the overall effects of fire on tree communities and the factors controlling their sensitivity in 29 sites that experienced multi-decadal alterations in fire frequencies in savanna and forest ecosystems across tropical and temperate regions. Fire had a strong overall effect on tree communities, with an average fire frequency (one fire every three years) reducing stem density by 48% and basal area by 53% after 50 years, relative to unburned plots. The largest changes occurred in savanna ecosystems and in sites with strong wet seasons or strong dry seasons, pointing to fire characteristics and species composition as important. Analyses of functional traits highlighted the impact of fire-driven changes in soil nutrients because frequent burning favoured trees with low biomass nitrogen and phosphorus content, and with more efficient nitrogen acquisition through ectomycorrhizal symbioses. Taken together, the response of trees to altered fire frequencies depends both on climatic and vegetation determinants of fire behaviour and tree growth, and the coupling between fire-driven nutrient losses and plant traits.

Rooting depth as a key woody functional trait in savannas.

Dimensions of tree root systems in savannas are poorly understood, despite being essential in resource acquisition and post-disturbance recovery. We studied tree rooting patterns in Southern African savannas to ask: how tree rooting strategies affected species responses to severe drought; and how potential rooting depths varied across gradients in soil texture and rainfall. First, detailed excavations of eight species in Kruger National Park suggest that the ratio of deep to shallow taproot diameters provides a reasonable proxy for potential rooting depth, facilitating extensive interspecific comparison. Detailed excavations also suggest that allocation to deep roots traded off with shallow lateral root investment, and that drought-sensitive species rooted more shallowly than drought-resistant ones. More broadly across 57 species in Southern Africa, potential rooting depths were phylogenetically constrained, with investment to deep roots evident among miombo Detarioids, consistent with results suggesting they green up before onset of seasonal rains. Soil substrate explained variation, with deeper roots on sandy, nutrient-poor soils relative to clayey, nutrient-rich ones. Although potential rooting depth decreased with increasing wet season length, mean annual rainfall had no systematic effect on rooting depth. Overall, our results suggest that rooting depth systematically structures the ecology of savanna trees. Further work examining other anatomical and physiological root traits should be a priority for understanding savanna responses to changing climate and disturbances.

Grasses continue to trump trees at soil carbon sequestration following herbivore exclusion in a semiarid African savanna.

Although studies have shown that mammalian herbivores often limit aboveground carbon storage in savannas, their effects on belowground soil carbon storage remain unclear. Using three sets of long-term, large herbivore exclosures with paired controls, we asked how almost two decades of herbivore removal from a semiarid savanna in Laikipia, Kenya affected aboveground (woody and grass) and belowground soil carbon sequestration, and determined the major source (C3 vs. C4 ) of belowground carbon sequestered in soils with and without herbivores present. Large herbivore exclusion, which included a diverse community of grazers, browsers, and mixed-feeding ungulates, resulted in significant increases in grass cover (~22%), woody basal area (~8 m2 /ha), and woody canopy cover (31%), translating to a ~8.5 t/ha increase in aboveground carbon over two decades. Herbivore exclusion also led to a 54% increase (20.5 t/ha) in total soil carbon to 30-cm depth, with ~71% of this derived from C4 grasses (vs. ~76% with herbivores present) despite substantial increases in woody cover. We attribute this continued high contribution of C4 grasses to soil C sequestration to the reduced offtake of grass biomass with herbivore exclusion together with the facilitative influence of open sparse woody canopies (e.g., Acacia spp.) on grass cover and productivity in this semiarid system.

Severe drought limits trees in a semi-arid savanna.

Increasingly frequent and severe droughts under climate change are expected to have major impacts on vegetation worldwide. However, research to date has focused on tree vulnerability to drought in forests. Less is known about trees and drought in savannas, where a sparse tree layer coexists with grass. These tree-grass interactions (often mediated by fire and herbivory) shape savanna tree ecology, and confound predictions of how strongly drought might affect trees. On the one hand, drought is physiologically stressful, which could harm trees and be exacerbated by herbivore impacts; on the other hand, trees adapted to semiarid savannas might be relatively drought tolerant, and the considerable impacts of drought on grass could even benefit trees via reduced grass competition and fire risk, especially in the year following a drought. Here, we sought to understand the net effects of severe drought on the savanna tree layer, and how fire and herbivory mediate these effects. We monitored tree growth, mortality, and community structure for 2 yr within existing long-term fire and herbivory experiments across a drought-severity contrast, following a major drought in Kruger National Park, South Africa. Overall, severe drought was a major stressor for trees. Tree mortality rates in most species increased by an order of magnitude in the year following drought, and slower growth rates for some persisted for 2 yr. At the community level, this translated into substantial decreases in tree densities. Herbivory and fire did little either to mitigate or exacerbate drought effects on trees, and overall, drought swamped effects of herbivory and fire that have otherwise been observed. However, species differed in their responses to drought, with some dominant encroaching species especially vulnerable. We suggest that increasing drought frequency and severity could drastically alter savanna vegetation by repeatedly killing off trees.

Defence strategies in African savanna trees.

Southern African savannas are commonly polarised into two broad types based on plant functional types and defences; infertile savannas dominated by broad-leaved trees typically defended by nitrogen-free secondary compounds and fertile savannas dominated by fine-leaved trees defended by structural defences. In this study, we use trait and other data from 15 wooded savanna sites in Southern Africa and ask if broad-leaved and fine-leaved species dominate on nutrient-poor and nutrient-rich soils, respectively. We then test if there is there any evidence for trade-offs in chemical (i.e., condensed tannins and total polyphenols) vs. structural defences on different soil types. We did not find strong evidence for a general divide in fine- vs. broad-leaved savannas according to soil fertility, nor for a simple trade-off between chemical and structural defences. Instead, we found savanna species to cluster into three broad defence strategies: species were high in leaf N and either (A) highly defended by spines and chemicals or (B) only structurally defended, or (C) low in leaf N and chemically defended. Finally, we tested for differences in browser utilisation between soil types and among plant defence strategies and found that browsing by meso-herbivores was higher on nutrient-rich soils and targeted species from groups A and B and avoided C, while browsing by elephants was mostly not affected by soil type or defence strategy. We propose a framework that can be used as a basis for asking strategic questions that will help improve our understanding of plant defences in savannas.

Carnivore stable carbon isotope niches reflect predator-prey size relationships in African savannas.

Predator-prey size relationships are among the most important patterns underlying the structure and function of ecological communities. Indeed, these relationships have already been shown to be important for understanding patterns of macroevolution and differential extinction in the terrestrial vertebrate fossil record. Stable isotope analysis (SIA) is a powerful remote approach to examining animal diets and paleodiets. The approach is based on the principle that isotope compositions of consumer tissues reflect those of their prey. In systems where resource isotope compositions are distributed along a body size gradient, SIA could be used to reconstruct predator-prey size relationships. We analyzed stable carbon isotope distributions amongst mammalian herbivores in extant and Plio-Pleistocene African savanna assemblages, and show that the range of δ13 C values among mammalian prey species (herbivores and rodents) increases with body mass (BM), because C4 plant feeding (essentially grazing) is more common among larger taxa. Consequently, δ13 C values of mammalian carnivores in these systems are related to species' BM, reflecting a higher average C4 prey component in the diets of larger-bodied carnivores. This pattern likely emerges because only the largest carnivores in these systems have regular access to the C4 prey base, whereas smaller carnivores do not. The δ13 C-BM relationship observed in mammalian carnivores is a potentially powerful approach for reconstructing and parameterizing predator-prey size relationships in contemporary and fossil savanna assemblages, and for interpreting how various behavioral, ecological and environmental factors influence prey size selection.

An overview of nitrogen cycling in a semiarid savanna: some implications for management and conservation in a large African park.

Nitrogen (N) is a major control on primary productivity and hence on the productivity and diversity of secondary producers and consumers. As such, ecosystem structure and function cannot be understood without a comprehensive understanding of N cycling and dynamics. This overview describes the factors that govern N distribution and dynamics and the consequences that variable N dynamics have for structure, function and thresholds of potential concern (TPCs) for management of a semiarid southern African savanna. We focus on the Kruger National Park (KNP), a relatively intact savanna, noted for its wide array of animal and plant species and a prized tourist destination. KNP's large size ensures integrity of most ecosystem processes and much can be learned about drivers of ecosystem structure and function using this park as a baseline. Our overview shows that large scale variability in substrates exists, but do not necessarily have predictable consequences for N cycling. The impact of major drivers such as fire is complex; at a landscape scale little differences in stocks and cycling were found, though at a smaller scale changes in woody cover can lead to concomitant changes in total N. Contrasting impacts of browsers and grazers on N turnover has been recorded. Due to the complexity of this ecosystem, we conclude that it will be complicated to draw up TPCs for most transformations and pools involved with the N cycle. However, we highlight in which cases the development of TPCs will be possible.

Frequent fire affects soil nitrogen and carbon in an African savanna by changing woody cover.

When tropical and sub-tropical ecosystems burn, considerable amounts of N present in the biomass fuel may be released. This ultimately results in a loss of fixed N to the atmosphere. It is often assumed that this volatilization loss of N with frequent fire will result in a reduction of plant-available N and total system N. By changing the amount of woody biomass fire may, however, also have indirect effects on N and C dynamics. Here we consider the effects of 50 years of frequent fire on total soil N and soil organic C (SOC) and total soil N in a mesic savanna in the Kruger National Park, South Africa. We also determine how changes in woody biomass may affect total soil N and SOC. We measured soil and fine root N and C concentrations as well as total soil N and SOC pools in four burning treatments, including fire exclusion, of a long-term fire experiment. Our results show that regardless of soil depth, fire treatment had no significant effect on total soil N and SOC. Our results also show that under trees total soil N and SOC concentrations of the surface soil increase, and pools of N and SOC increase to a depth of 7 cm. However, the extent to which soil N and C dynamics differed under canopies and away from canopies was dependent on fire treatment. Our results show that the effect of fire on soil N and C is mediated both through the indirect effect of changes in woody cover and the direct effects of fire (volatilization losses of nutrients). We suggest that woody thickening in this mesic savanna will have pronounced effects on long-term N and C dynamics.