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.

Reduced global fire activity due to human demography slows global warming by enhanced land carbon uptake.

Fire is an important climate-driven disturbance in terrestrial ecosystems, also modulated by human ignitions or fire suppression. Changes in fire emissions can feed back on the global carbon cycle, but whether the trajectories of changing fire activity will exacerbate or attenuate climate change is poorly understood. Here, we quantify fire dynamics under historical and future climate and human demography using a coupled global climate­fire­carbon cycle model that emulates 34 individual Earth system models (ESMs). Results are compared with counterfactual worlds, one with a constant preindustrial fire regime and another without fire. Although uncertainty in projected fire effects is large and depends on ESM, socioeconomic trajectory, and emissions scenario, we find that changes in human demography tend to suppress global fire activity, keeping more carbon within terrestrial ecosystems and attenuating warming. Globally, changes in fire have acted to warm climate throughout most of the 20th century. However, recent and predicted future reductions in fire activity may reverse this, enhancing land carbon uptake and corresponding to offsetting ∼5 to 10 y of global CO2 emissions at today's levels. This potentially reduces warming by up to 0.11 °C by 2100. We show that climate­carbon cycle feedbacks, as caused by changing fire regimes, are most effective at slowing global warming under lower emission scenarios. Our study highlights that ignitions and active and passive fire suppression can be as important in driving future fire regimes as changes in climate, although with some risk of more extreme fires regionally and with implications for other ecosystem functions in fire-dependent ecosystems.

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.

Grazing herbivores reduce herbaceous biomass and fire activity across African savannas.

Fire and herbivory interact to alter ecosystems and carbon cycling. In savannas, herbivores can reduce fire activity by removing grass biomass, but the size of these effects and what regulates them remain uncertain. To examine grazing effects on fuels and fire regimes across African savannas, we combined data from herbivore exclosure experiments with remotely sensed data on fire activity and herbivore density. We show that, broadly across African savannas, grazing herbivores substantially reduce both herbaceous biomass and fire activity. The size of these effects was strongly associated with grazing herbivore densities, and surprisingly, was mostly consistent across different environments. A one-zebra increase in herbivore biomass density (~100 kg/km2 of metabolic biomass) resulted in a ~53 kg/ha reduction in standing herbaceous biomass and a ~0.43 percentage point reduction in burned area. Our results indicate that fire models can be improved by incorporating grazing effects on grass biomass.

Carbon isotope trends across a century of herbarium specimens suggest CO2 fertilization of C4 grasses.

Increasing atmospheric CO2 is changing the dynamics of tropical savanna vegetation. C3 trees and grasses are known to experience CO2 fertilization, whereas responses to CO2 by C4 grasses are more ambiguous. Here, we sample stable carbon isotope trends in herbarium collections of South African C4 and C3 grasses to reconstruct 13C discrimination. We found that C3 grasses showed no trends in 13C discrimination over the past century but that C4 grasses increased their 13C discrimination through time, especially since 1950. These changes were most strongly linked to changes in atmospheric CO2 rather than to trends in rainfall climatology or temperature. Combined with previously published evidence that grass biomass has increased in C4-dominated savannas, these trends suggest that increasing water-use efficiency due to CO2 fertilization may be changing C4 plant-water relations. CO2 fertilization of C4 grasses may thus be a neglected pathway for anthropogenic global change in tropical savanna ecosystems.

Fire frequency drives decadal changes in soil carbon and nitrogen and ecosystem productivity.

Fire frequency is changing globally and is projected to affect the global carbon cycle and climate. However, uncertainty about how ecosystems respond to decadal changes in fire frequency makes it difficult to predict the effects of altered fire regimes on the carbon cycle; for instance, we do not fully understand the long-term effects of fire on soil carbon and nutrient storage, or whether fire-driven nutrient losses limit plant productivity. Here we analyse data from 48 sites in savanna grasslands, broadleaf forests and needleleaf forests spanning up to 65 years, during which time the frequency of fires was altered at each site. We find that frequently burned plots experienced a decline in surface soil carbon and nitrogen that was non-saturating through time, having 36 per cent (±13 per cent) less carbon and 38 per cent (±16 per cent) less nitrogen after 64 years than plots that were protected from fire. Fire-driven carbon and nitrogen losses were substantial in savanna grasslands and broadleaf forests, but not in temperate and boreal needleleaf forests. We also observe comparable soil carbon and nitrogen losses in an independent field dataset and in dynamic model simulations of global vegetation. The model study predicts that the long-term losses of soil nitrogen that result from more frequent burning may in turn decrease the carbon that is sequestered by net primary productivity by about 20 per cent of the total carbon that is emitted from burning biomass over the same period. Furthermore, we estimate that the effects of changes in fire frequency on ecosystem carbon storage may be 30 per cent too low if they do not include multidecadal changes in soil carbon, especially in drier savanna grasslands. Future changes in fire frequency may shift ecosystem carbon storage by changing soil carbon pools and nitrogen limitations on plant growth, altering the carbon sink capacity of frequently burning savanna grasslands and broadleaf forests.

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.

Seasonal strategies differ between tropical and extratropical herbivores.

Seasonal diet shifts and migration are key components of large herbivore population dynamics, but we lack a systematic understanding of how these behaviours are distributed on a macroecological scale. The prevalence of seasonal strategies is likely related to herbivore body size and feeding guild, and may also be influenced by properties of the environment, such as soil nutrient availability and climate seasonality. We evaluated the distribution of seasonal dietary shifts and migration across large-bodied mammalian herbivores and determined how these behaviours related to diet, body size and environment. We found that herbivore strategies were consistently correlated with their traits: seasonal diet shifts were most prevalent among mixed feeding herbivores and migration among grazers and larger herbivores. Seasonality also played a role, particularly for migration, which was more common at higher latitudes. Both dietary shifts and migration were more widespread among extratropical herbivores, which also exhibited more intermediate diets and body sizes. Our findings suggest that strong seasonality in extratropical systems imposes pressure on herbivores, necessitating widespread behavioural responses to navigate seasonal resource bottlenecks. It follows that tropical and extratropical herbivores may have divergent responses to global change, with intensifying herbivore pressure in extratropical systems contrasting with diminishing herbivore pressure in tropical systems.

Spatial patterning among savanna trees in high-resolution, spatially extensive data.

In savannas, predicting how vegetation varies is a longstanding challenge. Spatial patterning in vegetation may structure that variability, mediated by spatial interactions, including competition and facilitation. Here, we use unique high-resolution, spatially extensive data of tree distributions in an African savanna, derived from airborne Light Detection and Ranging (LiDAR), to examine tree-clustering patterns. We show that tree cluster sizes were governed by power laws over two to three orders of magnitude in spatial scale and that the parameters on their distributions were invariant with respect to underlying environment. Concluding that some universal process governs spatial patterns in tree distributions may be premature. However, we can say that, although the tree layer may look unpredictable locally, at scales relevant to prediction in, e.g., global vegetation models, vegetation is instead strongly structured by regular statistical distributions.

Disease and fire interact to influence transitions between savanna-forest ecosystems over a multi-decadal experiment.

Global change is shifting disturbance regimes that may rapidly change ecosystems, sometimes causing ecosystems to shift between states. Interactions between disturbances such as fire and disease could have especially severe effects, but experimental tests of multi-decadal changes in disturbance regimes are rare. Here, we surveyed vegetation for 35 years in a 54-year fire frequency experiment in a temperate oak savanna-forest ecotone that experienced a recent outbreak of oak wilt. Different fire regimes determined whether plots were savanna or forest by regulating tree abundance (r2  = 0.70), but disease rapidly reversed the effect of fire exclusion, increasing mortality by 765% in unburned forests, but causing relatively minor changes in frequently burned savannas. Model simulations demonstrated that disease caused unburned forests to transition towards a unique woodland that was prone to transition to savanna if fire was reintroduced. Consequently, disease-fire interactions could shift ecosystem resilience and biome boundaries as pathogen distributions change.

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.

Dispersal Increases the Resilience of Tropical Savanna and Forest Distributions.

Global change may induce changes in savanna and forest distributions, but the dynamics of these changes remain unclear. Classical biome theory suggests that climate is predictive of biome distributions, such that shifts will be continuous and reversible. This view, however, cannot explain the overlap in the climatic ranges of tropical biomes, which some argue may result from fire-vegetation feedbacks, maintaining savanna and forest as bistable states. Under this view, biome shifts are argued to be discontinuous and irreversible. Mean-field bistable models, however, are also limited, as they cannot reproduce the spatial aggregation of biomes. Here we suggest that both models ignore spatial processes, such as dispersal, which may be important when savanna and forest abut. We examine the contributions of dispersal to determining biome distributions using a 2D reaction-diffusion model, comparing results qualitatively to empirical savanna and forest distributions in sub-Saharan Africa. We find that the diffusion model resolves both the aforementioned limitations of biome models. First, local dispersive spatial interactions, with an underlying precipitation gradient, can reproduce the spatial aggregation of biomes with a stable savanna-forest boundary. Second, the boundary is determined not only by the amount of precipitation but also by the geometrical shape of the precipitation contours. These geometrical effects arise from continental-scale source-sink dynamics, which reproduce the mismatch between biome and climate. Dynamically, the spatial model predicts that dispersal may increase the resilience of tropical biome in response to global change: the boundary continuously tracks climate, recovering following disturbances, unless the remnant biome patches are too small.

Determinants of tree cover in tropical floodplains.

Tree cover differentiates forests from savannas and grasslands. In tropical floodplains, factors differentiating these systems are poorly known, even though floodplains cover 10% of the tropical landmass. Seasonal inundation potentially presents trees with both challenges (soil anoxia) and benefits (moisture and nutrient deposition), the relative importance of which may depend on ecological context, e.g. if floods alleviate water stress more in more arid ecosystems. Here, we use remotely sensed data across 13 large tropical and sub-tropical floodplain ecosystems on five continents to show that climatic water balance (i.e. precipitation-potential evapotranspiration) strongly increases floodplain tree cover in interaction with flooding, fire and topography. As predicted, flooding increases tree cover in more arid floodplains, but decreases tree cover in climatically wetter ones. As in uplands, frequent fire reduced tree cover, particularly in wet regions, but-in contrast with uplands-lower elevation and sandier soils decreased tree cover. Our results suggest that predicting the impacts of changing climate, land use and hydrology on floodplain ecosystems depends on considering climate-disturbance interactions. While outright wetland conversion proceeds globally, additional anthropogenic activities, including alteration of fire frequencies and dam construction, will also shift floodplain tree cover, especially in wet climates.

Palaeo-trajectories of forest savannization in the southern Congo.

Tropical savannah and forest are thought to represent alternative stable states in ecosystem structure in some climates. The implication is that biomes are maintained by positive feedbacks, e.g. with fire, and that historical distributions could play a role in determining modern ones. In this context, climate alone does not govern transitions between biomes, and understanding the causes and pathways of such transitions becomes crucial. Here, we use a multi-proxy analysis of a 2000-year core to evaluate modes of transition in vegetation structure and fire regimes. We demonstrate a first transition ca 1540 BP, when a cyclic fire regime entered a forested landscape, eventually resulting, by ca 1060 BP, in a transition to a more open savannah-like or mosaicked structure. This pattern may parallel currently accelerating fire regimes in tropical forests suggesting that fires can savannize forests, but perhaps more slowly than feared. Finally, ca 540 BP, a drought combined with anthropogenic influences resulted in a conclusive transition to savannah, probably resembling the modern landscape in the region. We show here that fire interacted with drought to transition forest to savannah, suggesting that disturbance by fire can be a major driver of biome change.

Prediction and scale in savanna ecosystems.

Contents Summary 52 I. Introduction 52 II. Determinants of savanna vegetation structure 53 III. Are trees in savannas really more heterogeneous? 53 IV. Are trees in savannas a 'slow' variable? 54 V. Are trees in savannas spatially patterned? 55 VI. Conclusions 55 Acknowledgements 55 References 56 SUMMARY: Savannas are highly variable systems, and predicting variation, especially in the tree layer, represents a major unresolved challenge for forecasting biosphere responses to global change. Prediction to date has focused on disentangling interactions between resource limitation and chronic disturbances to identify what determines local savanna vegetation heterogeneity. By focusing at too fine a scale, this approach overlooks: sample size limitation arising from sparse tree distributions; stochasticity in demographic and environmental processes that is preserved as heterogeneity among tree populations with slow dynamics; and spatial self-organization. Renewed focus on large (1-50 ha) permanent plots and on spatial patterns of tree-layer variability at even larger landscape spatial scales (≥1000s of ha) promises to resolve these limitations, consistent with the goal of predicting large-scale biosphere responses to global change.

Soil texture mediates tree responses to rainfall intensity in African savannas.

Rainfall variability is a major determinant of soil moisture, but its influence on vegetation structure has been challenging to generalize. This presents a major source of uncertainty in predicting vegetation responses to potentially widespread shifts in rainfall frequency and intensity. In savannas, where trees and grasses coexist, conflicting lines of evidence have suggested, variously, that tree cover can either increase or decrease in response to less frequent, more intense rainfall. Here, we use remote sensing products and continent-wide soil maps for sub-Saharan Africa to analyze how soil texture and fire mediate the response of savanna tree cover to rainfall climatology. Tree cover increased with mean wet-season rainfall and decreased with fire frequency, consistent with previous analyses. However, responses to rainfall intensity varied: tree cover dramatically decreased with rainfall intensity on clayey soils, at high rainfall, and with rainfall spread over longer wet seasons; conversely, on sandy soils, at low rainfall, and with shorter wet seasons, tree cover instead increased with rainfall intensity. Tree cover responses to rainfall climatology depend on soil texture, accounting for substantial variation in tree cover across African savannas. Differences in underlying soils may lead to divergent responses of savannas to global change.

Soils and fire jointly determine vegetation structure in an African savanna.

Savanna vegetation is variable, and predicting how water, nutrients, and chronic disturbances interact to determine vegetation structure in savannas represents a challenge. Here, we examined in situ interactions among rainfall, soils, grasses, fire, and elephants that determine tree layer responses to resource gradients in Kruger National Park in South Africa, using 363 long-term monitoring sites throughout the park. Grass biomass increased with rainfall and on nutrient-rich clay soils. Fire frequency, too, increased with rainfall. Conversely, tree density was greater on sandier soils, where water infiltrates more readily, and in areas where the maximum interval between fires was longer, irrespective of average fire frequency. Elephant density responded positively to tree density, but did not contribute significantly to decreasing tree density. Savanna vegetation structure was reasonably predictable, via a combination of rainfall (favoring grasses), soil (sandy soils favoring trees), and fire (limiting trees until a longer interval between fires allows them to establish). Explicit consideration of bottom-up and top-down interactions may thus contribute to a predictive understanding of savanna vegetation heterogeneity.

Aridity, not fire, favors nitrogen-fixing plants across tropical savanna and forest biomes.

Tropical savannas are hypothesized to be hot spots of nitrogen-fixer diversity and activity because of the high disturbance and low nitrogen characteristic of savanna landscapes. Here we compare the abundances of nitrogen-fixing and non-fixing trees in both tropical savannas and tropical forests under climatically equivalent conditions, using plant inventory studies across 566 plots in South America and Africa. A single factor, aridity, explained 19-54% of the variance in fixer abundance, and unexpectedly was more important than fire frequency, biome, and continent. Nitrogen fixers were more abundant in arid environments; as a result, African savannas, which tend to be drier, were richer in nitrogen fixers than South American savannas. Fixer abundance converged on similar levels in forests in both continents. We conclude that climate plays a greater role than fire in determining the distribution of nitrogen fixers across tropical savanna and forest biomes.

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 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.

Evolution of human-driven fire regimes in Africa.

Human ability to manipulate fire and the landscape has increased over evolutionary time, but the impact of this on fire regimes and consequences for biodiversity and biogeochemistry are hotly debated. Reconstructing historical changes in human-derived fire regimes empirically is challenging, but information is available on the timing of key human innovations and on current human impacts on fire; here we incorporate this knowledge into a spatially explicit fire propagation model. We explore how changes in population density, the ability to create fire, and the expansion of agropastoralism altered the extent and seasonal distribution of fire as modern humans arose and spread through Africa. Much emphasis has been placed on the positive effect of population density on ignition frequency, but our model suggests this is less important than changes in fire spread and connectivity that would have occurred as humans learned to light fires in the dry season and to transform the landscape through grazing and cultivation. Different landscapes show different limitations; we show that substantial human impacts on burned area would only have started ~4,000 B.P. in open landscapes, whereas they could have altered fire regimes in closed/dissected landscapes by ~40,000 B.P. Dry season fires have been the norm for the past 200-300 ky across all landscapes. The annual area burned in Africa probably peaked between 4 and 40 kya. These results agree with recent paleocarbon studies that suggest that the biomass burned today is less than in the recent past in subtropical countries.