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.

Fire facilitates ground layer plant diversity in a Miombo ecosystem.

BACKGROUND AND AIMS: Little is known about the response of ground layer plant communities to fire in Miombo ecosystems, which is a global blind spot of ecological understanding. We aimed: (1) to assess the impact of three experimentally imposed fire treatments on ground layer species composition and compare it with patterns observed for trees; and (2) to analyse the effect of fire treatments on species richness to assess how responses differ among plant functional groups. METHODS: At a 60-year-long fire experiment in Zambia, we quantified the richness and diversity of ground layer plants in terms of taxa and functional groups across three experimental fire treatments of late dry-season fire, early dry-season fire and fire exclusion. Data were collected in five repeat surveys from the onset of the wet season to the early dry season. KEY RESULTS: Of the 140 ground layer species recorded across the three treatments, fire-maintained treatments contributed most of the richness and diversity, with the least number of unique species found in the no-fire treatment. The early-fire treatment was more similar in composition to the no-fire treatment than to the late-fire treatment. C4 grass and geoxyle richness were highest in the late-fire treatment, and there were no shared sedge species between the late-fire and other treatments. At a plot level, the average richness in the late-fire treatment was twice that of the fire exclusion treatment. CONCLUSIONS: Heterogeneity in fire seasonality and intensity supports diversity of a unique flora by providing a diversity of local environments. African ecosystems face rapid expansion of land- and fire-management schemes for carbon offsetting and sequestration. We demonstrate that analyses of the impacts of such schemes predicated on the tree flora alone are highly likely to underestimate impacts on biodiversity. A research priority must be a new understanding of the Miombo ground layer flora integrated into policy and land management.

Resprouting grasses are associated with less frequent fire than seeders.

Plant populations persist under recurrent fire via resprouting from surviving tissues (resprouters) or seedling recruitment (seeders). Woody species are inherently slow maturing, meaning that seeders are confined to infrequent fire regimes. However, for grasses, which mature faster, the relationships between persistence strategy and fire regime remain unknown. Globally, we analysed associations between fire regimes experienced by hundreds of grass species and their persistence strategy, within a phylogenetic context. We also tested whether persistence strategies are associated with morphological and physiological traits. Resprouters were associated with less frequent fire than seeders. Whilst modal fire frequencies were similar (fire return interval of 4-6 yr), seeders were restricted to regions with more frequent fire than resprouters, suggesting that greater competition with long-lived resprouters restricts seeder recruitment and survival when fire is rare. Resprouting was associated with lower leaf N, higher C:N ratios and the presence of belowground buds, but was unrelated to photosynthetic pathway. Differences between the life histories of grasses and woody species led to a contrasting prevalence of seeders and resprouters in relation to fire frequency. Rapid sexual maturation in grasses means that seeder distributions, relative to fire regime, are determined by competitive ability and recruitment, rather than time to reproductive maturity.

Global ecosystems and fire: Multi-model assessment of fire-induced tree-cover and carbon storage reduction.

In this study, we use simulations from seven global vegetation models to provide the first multi-model estimate of fire impacts on global tree cover and the carbon cycle under current climate and anthropogenic land use conditions, averaged for the years 2001-2012. Fire globally reduces the tree covered area and vegetation carbon storage by 10%. Regionally, the effects are much stronger, up to 20% for certain latitudinal bands, and 17% in savanna regions. Global fire effects on total carbon storage and carbon turnover times are lower with the effect on gross primary productivity (GPP) close to 0. We find the strongest impacts of fire in savanna regions. Climatic conditions in regions with the highest burned area differ from regions with highest absolute fire impact, which are characterized by higher precipitation. Our estimates of fire-induced vegetation change are lower than previous studies. We attribute these differences to different definitions of vegetation change and effects of anthropogenic land use, which were not considered in previous studies and decreases the impact of fire on tree cover. Accounting for fires significantly improves the spatial patterns of simulated tree cover, which demonstrates the need to represent fire in dynamic vegetation models. Based upon comparisons between models and observations, process understanding and representation in models, we assess a higher confidence in the fire impact on tree cover and vegetation carbon compared to GPP, total carbon storage and turnover times. We have higher confidence in the spatial patterns compared to the global totals of the simulated fire impact. As we used an ensemble of state-of-the-art fire models, including effects of land use and the ensemble median or mean compares better to observational datasets than any individual model, we consider the here presented results to be the current best estimate of global fire effects on ecosystems.

A unified framework for plant life-history strategies shaped by fire and herbivory.

Fire and herbivory both remove aboveground biomass. Environmental factors determine the type and intensity of these consumers globally, but the traits of plants can also alter their propensity to burn and the degree to which they are eaten. To understand plant life-history strategies associated with fire and herbivory we need to describe both response and effect functional traits, and how they sort within communities, along resource gradients, and across evolutionary timescales. Fire and herbivore functional traits are generally considered separately, but there are advances made in understanding fire that relate to herbivory, and vice versa. Moreover, fire and herbivory interact: the presence of one consumer affects the type and intensity of the other. Here, we present a unifying conceptual framework to understand plant strategies that enable tolerance and persistence to fire and herbivory. Using grasses as an example, we discuss how flammability and fire tolerance, palatability, and grazing tolerance traits might organize themselves in ecosystems exposed to these consumers, and how these traits might have evolved with reference to other strong selective processes, like aridity. Our framework can be used to predict both the diversity of life-history strategies and plant species diversity under different consumer regimes.

Anthropogenic modifications to fire regimes in the wider Serengeti-Mara ecosystem.

Fire is a key driver in savannah systems and widely used as a land management tool. Intensifying human land uses are leading to rapid changes in the fire regimes, with consequences for ecosystem functioning and composition. We undertake a novel analysis describing spatial patterns in the fire regime of the Serengeti-Mara ecosystem, document multidecadal temporal changes and investigate the factors underlying these patterns. We used MODIS active fire and burned area products from 2001 to 2014 to identify individual fires; summarizing four characteristics for each detected fire: size, ignition date, time since last fire and radiative power. Using satellite imagery, we estimated the rate of change in the density of livestock bomas as a proxy for livestock density. We used these metrics to model drivers of variation in the four fire characteristics, as well as total number of fires and total area burned. Fires in the Serengeti-Mara show high spatial variability-with number of fires and ignition date mirroring mean annual precipitation. The short-term effect of rainfall decreases fire size and intensity but cumulative rainfall over several years leads to increased standing grass biomass and fuel loads, and, therefore, in larger and hotter fires. Our study reveals dramatic changes over time, with a reduction in total number of fires and total area burned, to the point where some areas now experience virtually no fire. We suggest that increasing livestock numbers are driving this decline, presumably by inhibiting fire spread. These temporal patterns are part of a global decline in total area burned, especially in savannahs, and we caution that ecosystem functioning may have been compromised. Land managers and policy formulators need to factor in rapid fire regime modifications to achieve management objectives and maintain the ecological function of savannah ecosystems.

Pyrodiversity interacts with rainfall to increase bird and mammal richness in African savannas.

Fire is a fundamental process in savannas and is widely used for management. Pyrodiversity, variation in local fire characteristics, has been proposed as a driver of biodiversity although empirical evidence is equivocal. Using a new measure of pyrodiversity (Hempson et al.), we undertook the first continent-wide assessment of how pyrodiversity affects biodiversity in protected areas across African savannas. The influence of pyrodiversity on bird and mammal species richness varied with rainfall: strongest support for a positive effect occurred in wet savannas (> 650 mm/year), where species richness increased by 27% for mammals and 40% for birds in the most pyrodiverse regions. Range-restricted birds were most increased by pyrodiversity, suggesting the diversity of fire regimes increases the availability of rare niches. Our findings are significant because they explain the conflicting results found in previous studies of savannas. We argue that managing savanna landscapes to increase pyrodiversity is especially important in wet savannas.

Management impacts on fire occurrence: A comparison of fire regimes of African and South American tropical savannas in different protected areas.

Humans can alter fire dynamics in grassland systems by changing fire frequency, fire seasonality and fuel conditions. These changes have effects on vegetation structure and recovery, species composition, and ecosystem function. Understanding how human management can affect fire regimes is vital to detect potential changes in the resilience of plant communities, and to predict vegetation responses to human interventions. We evaluated the fire regimes of two recently protected areas in Madagascar (Ibity and Itremo NPA) and one in Brazil (Serra do Cipó NP) before and after livestock exclusion and fire suppression policies. We compare the pre- and post-management fire history in these areas and analyze differences in terms of total annual burned area, density of ignitions, burn scar size distribution, fire return period and seasonal fire distribution. More than 90% of total park areas were burned at least once during the studied period, for all parks. We observed a significant reduction in the number of ignitions for Ibity NPA and Serra do Cipó NP after livestock exclusion and active fire suppression, but no significant change in total burned area for each protected area. We also observed a seasonal shift in burning, with fires happening later in the fire season (October-November) after management intervention. However, the protected areas in Madagascar had shorter fire return intervals (3.23 and 1.82 years) than those in Brazil (7.91 years). Our results demonstrate that fire exclusion is unattainable, and probably unwarranted in tropical grassland conservation areas, but show how human intervention in fire and vegetation patterns can alter various aspects of the fire regimes. This information can help with formulating realistic and effective fire management policies in these valuable conservation areas.

Defining pyromes and global syndromes of fire regimes.

Fire is a ubiquitous component of the Earth system that is poorly understood. To date, a global-scale understanding of fire is largely limited to the annual extent of burning as detected by satellites. This is problematic because fire is multidimensional, and focus on a single metric belies its complexity and importance within the Earth system. To address this, we identified five key characteristics of fire regimes--size, frequency, intensity, season, and extent--and combined new and existing global datasets to represent each. We assessed how these global fire regime characteristics are related to patterns of climate, vegetation (biomes), and human activity. Cross-correlations demonstrate that only certain combinations of fire characteristics are possible, reflecting fundamental constraints in the types of fire regimes that can exist. A Bayesian clustering algorithm identified five global syndromes of fire regimes, or pyromes. Four pyromes represent distinctions between crown, litter, and grass-fueled fires, and the relationship of these to biomes and climate are not deterministic. Pyromes were partially discriminated on the basis of available moisture and rainfall seasonality. Human impacts also affected pyromes and are globally apparent as the driver of a fifth and unique pyrome that represents human-engineered modifications to fire characteristics. Differing biomes and climates may be represented within the same pyrome, implying that pathways of change in future fire regimes in response to changes in climate and human activity may be difficult to predict.

Fire ecology of C3 and C4 grasses depends on evolutionary history and frequency of burning but not photosynthetic type.

Grasses using the C4 photosynthetic pathway dominate frequently burned savannas, where the pathway is hypothesized to be adaptive. However, independent C4 lineages also sort among different fire environments. Adaptations to fire may thus depend on evolutionary history, which could be as important as the possession of the C4 photosynthetic pathway for life in these environments. Here, using a comparative pot experiment and controlled burn, we examined C3 and C4 grasses belonging to four lineages from the same regional flora, and asked the following questions: Do lineages differ in their responses to fire, are responses consistent between photosynthetic types, and are responses related to fire frequency in natural habitats? We found that in the C4 Andropogoneae lineage, frost killed a large proportion of aboveground biomass and produced a large dry fuel load, which meant that only a small fraction of the living tissue was lost in the fire. C3 species from the Paniceae and Danthonioideae lineages generated smaller fuel loads and lost more living biomass, while species from the C4 lineage Aristida generated the smallest fuel loads and lost the most living tissue. Regrowth after the fire was more rapid and complete in the C4 Andropogoneae and C3 Paniceae, but incomplete and slower in the C3 Danthonioideae and C4 Aristida. Rapid recovery was associated with high photosynthetic rates, high specific leaf area, delayed flowering, and frequent fires in natural habitats. Results demonstrated that phylogenetic lineage was more important than photosynthetic type in determining the fire response of these grasses and that fire responses were related to the frequency that natural habitats burned.

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.

Increasing temperatures can improve seedling establishment in arid-adapted savanna trees.

Plant species are shifting their ranges in response to global climate change, thus intensifying the need to predict such changes accurately. As the environmental requirements controlling plant distribution act differently at each developmental stage, there is a need to acquire a demographic-specific understanding of the factors which determine these distributions. Here we investigated the germination niche of two common savanna species Acacia nigrescens and Colophospermum mopane, with the aims to disentangle the direct and indirect effects of temperature on seed germination and establishment and to explore the impact of higher temperatures on the establishment success of savanna trees. Under laboratory conditions, we used thermal gradient plates to determine the thermal germination niche of both species, and a water stress experiment was conducted on C. mopane to account for water-temperature interactions. Using these data we parameterised a soil-moisture model to determine germination and establishment success under field conditions at current and future temperatures (+4 °C). Based on this model, higher future temperatures will not limit germination directly, but they will reduce the number of germination events by reducing the time window of suitable available soil water. Conversely, warmer conditions will accelerate the rate of radicle extension and increase the frequency of seedling establishment events. An additional advantage of higher temperatures is that fewer seeds will germinate, resulting in slower seed bank depletion when successful seedling establishment events do occur.

Fire-mediated effects on polyploid biology.

Cold temperatures have been posited as a key driver of polyploidy (possession of multiple chromosome sets). However, high temperatures associated with fire, and the indirect impact of post-fire environments in polypoid formation and establishment deserve more attention for a comprehensive understanding of polyploid ecology, evolution, and current distributions.

Savanna fire regimes depend on grass trait diversity.

Grasses fuel most fires on Earth and strongly influence local fire behaviour through traits that determine how flammable they are. Therefore, grass communities that differ in their species and trait compositions give rise to significant spatial variation in savanna fire regimes across the world, which cannot be otherwise explained. Likewise, fire regimes are continuously modified by alterations to savanna grass community traits, through species introductions and climatic changes. However, current representation of grassy fuels in global fire models misses important variation and therefore limits predictive power. The inclusion of grass trait diversity in models, using remotely sensed trait proxies, for example, will greatly improve our ability to understand and project savanna fires and their roles in the Earth system.

Ancient grasslands guide ambitious goals in grassland restoration.

Grasslands, which constitute almost 40% of the terrestrial biosphere, provide habitat for a great diversity of animals and plants and contribute to the livelihoods of more than 1 billion people worldwide. Whereas the destruction and degradation of grasslands can occur rapidly, recent work indicates that complete recovery of biodiversity and essential functions occurs slowly or not at all. Grassland restoration-interventions to speed or guide this recovery-has received less attention than restoration of forested ecosystems, often due to the prevailing assumption that grasslands are recently formed habitats that can reassemble quickly. Viewing grassland restoration as long-term assembly toward old-growth endpoints, with appreciation of feedbacks and threshold shifts, will be crucial for recognizing when and how restoration can guide recovery of this globally important ecosystem.

Grazing lawns and overgrazing in frequently grazed grass communities.

Frequent grazing can establish high forage value grazing lawns supporting high grazer densities, but can also produce overgrazed grass communities with unpalatable or low grass basal cover, supporting few grazers. Attempts to create grazing lawns via concentrated grazing, with a goal to increase grazer numbers, are thus risky without knowing how environmental conditions influence the likelihood of each outcome. We collected grass species and trait data from 33 frequently grazed grass communities across eastern South Africa (28 sites) and the Serengeti National Park, Tanzania (five sites), covering wide rainfall (336-987 mm year-1) and soil (e.g., 44%-93% sand) gradients. We identified four grass growth forms using hierarchical clustering on principal components analyses of trait data and assessed trait-environment and growth form-environment relationships using fourth corner and principal components analyses. We distinguished two palatable grass growth forms that both attract yet resist grazers and comprise grazing lawns: (1) "lateral attractors" that spread vegetatively via stolons and rhizomes, and (2) "tufted attractors" that form isolated tufts and may have alternate tall growth forms. By contrast, (3) tough, upright, tufted "resisters," and (4) "avoiders" with sparse architectures or that grow appressed to the soil surface, are of little forage value and avoided by grazers. Grazing lawns occurred across a wide range of conditions, typically comprising lateral attractor grasses in drier, sandy environments, and tufted attractor grasses in wetter, low-sand environments. Resisters occurred on clay-rich soils in mesic areas, while avoiders were widespread but scarce. While grazing lawns can be established under most conditions, monitoring their composition and cover is important, as the potential for overgrazing seems as widely relevant. Tufted attractor-dominated lawns appear somewhat more vulnerable to degradation than lateral attractor-dominated lawns. Increased avoider and resister abundance both reduce forage value, although resisters may provide better soil protection.

Deciphering the distribution of the savanna biome.

• We aimed to identify the limits of savanna across Africa, Australia and South America. We based our investigation on the rich history of hypotheses previously examined: that the limits of savanna are variously determined by rainfall, rainfall seasonality, soil fertility and disturbance. • We categorized vegetation on all continents as 'savanna' (open habitats with a C(4) grass layer) or 'not-savanna' (closed habitats with no C(4) grass layer) and used a combination of statistical approaches to examine how the presence of savanna varied as a function of five environmental correlates. • The presence of savanna is constrained by effective rainfall and rainfall seasonality. Soil fertility is regionally important, although the direction of its effect changes relative to rainfall. We identified three continental divergences in the limits of savanna that could not be explained by environment. • Climate and soils do not have a deterministic effect on the distribution of savanna. Over the range of savanna, some proportion of the land is always 'not-savanna'. We reconciled previous contradictory views of savanna limits by developing a new conceptual framework for understanding these limits by categorizing environmental factors into whether they had a positive or negative effect on woody growth and the frequency of disturbance.

Tree cover in sub-Saharan Africa: rainfall and fire constrain forest and savanna as alternative stable states.

Savannas are known as ecosystems with tree cover below climate-defined equilibrium values. However, a predictive framework for understanding constraints on tree cover is lacking. We present (a) a spatially extensive analysis of tree cover and fire distribution in sub-Saharan Africa, and (b) a model, based on empirical results, demonstrating that savanna and forest may be alternative stable states in parts of Africa, with implications for understanding savanna distributions. Tree cover does not increase continuously with rainfall, but rather is constrained to low (<50%, "savanna") or high tree cover (>75%, "forest"). Intermediate tree cover rarely occurs. Fire, which prevents trees from establishing, differentiates high and low tree cover, especially in areas with rainfall between 1000 mm and 2000 mm. Fire is less important at low rainfall (<1000 mm), where rainfall limits tree cover, and at high rainfall (>2000 mm), where fire is rare. This pattern suggests that complex interactions between climate and disturbance produce emergent alternative states in tree cover. The relationship between tree cover and fire was incorporated into a dynamic model including grass, savanna tree saplings, and savanna trees. Only recruitment from sapling to adult tree varied depending on the amount of grass in the system. Based on our empirical analysis and previous work, fires spread only at tree cover of 40% or less, producing a sigmoidal fire probability distribution as a function of grass cover and therefore a sigmoidal sapling to tree recruitment function. This model demonstrates that, given relatively conservative and empirically supported assumptions about the establishment of trees in savannas, alternative stable states for the same set of environmental conditions (i.e., model parameters) are possible via a fire feedback mechanism. Integrating alternative stable state dynamics into models of biome distributions could improve our ability to predict changes in biome distributions and in carbon storage under climate and global change scenarios.