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.

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.

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.

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.

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.

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.

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.

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.

Fire and biodiversity in the Anthropocene.

Fire has been a source of global biodiversity for millions of years. However, interactions with anthropogenic drivers such as climate change, land use, and invasive species are changing the nature of fire activity and its impacts. We review how such changes are threatening species with extinction and transforming terrestrial ecosystems. Conservation of Earth's biological diversity will be achieved only by recognizing and responding to the critical role of fire. In the Anthropocene, this requires that conservation planning explicitly includes the combined effects of human activities and fire regimes. Improved forecasts for biodiversity must also integrate the connections among people, fire, and ecosystems. Such integration provides an opportunity for new actions that could revolutionize how society sustains biodiversity in a time of changing fire activity.

What drives grassland-forest boundaries? Assessing fire and frost effects on tree seedling survival and architecture.

Fire and frost represent two major hurdles for the persistence of trees in open grassy biomes and have both been proposed as drivers of grassland-forest boundaries in Africa.We assess the response of young tree seedlings, which represent a vulnerable stage in tree recruitment, to traumatic fire and frost disturbances.In a greenhouse experiment, we investigated how seedling traits predicted survival and resprouting ability in response to fire versus frost; we characterized survival strategies of seedlings in response to the two disturbances, and we documented how the architecture of surviving seedlings is affected by fire versus frost injury.Survival rates were similar under both treatments. However, different species displayed different levels of sensitivity to fire and frost. Seedling survival was higher for older seedlings and seedlings with more basal leaves. Survivors of a fire event lost more biomass than the survivors of a frost event. However, the architecture of recovered fire- and frost-treated seedlings was mostly similar. Seedlings that recovered from fire and frost treatments were often shorter than those that had not been exposed to any disturbance, with multiple thin branches, which may increase vulnerability to the next frost or fire event. Synthesis. Fire caused more severe aboveground damage compared with a single frost event, suggesting that fire is an important driver of tree distribution in these open grassland systems. However, the impact of repeated frost events may be equally severe and needs to be investigated. Also, woody species composition may be influenced by phenomena that affect the timing and frequency of seedling exposure to damage, as mortality was found to be dependent on seedling age. Therefore, changes in fire regime and climate are likely to result in changes in the composition and the structure of the woody components of these systems.

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.

Comment on "The global tree restoration potential".

Bastin et al's estimate (Reports, 5 July 2019, p. 76) that tree planting for climate change mitigation could sequester 205 gigatonnes of carbon is approximately five times too large. Their analysis inflated soil organic carbon gains, failed to safeguard against warming from trees at high latitudes and elevations, and considered afforestation of savannas, grasslands, and shrublands to be restoration.

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.

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.

Alternate Grassy Ecosystem States Are Determined by Palatability-Flammability Trade-Offs.

Fire and mammalian grazers both consume grasses, and feedbacks between grass species, their functional traits, and consumers have profound effects on grassy ecosystem structure worldwide, such that savanna and grassland states determined by fire or grazing can be considered alternate states. These parallel savanna-forest alternate states, which likewise have myriad cascading ecosystem impacts.

Can trophic rewilding reduce the impact of fire in a more flammable world?

Large vertebrates affect fire regimes in several ways: by consuming plant matter that would otherwise accumulate as fuel; by controlling and varying the density of vegetation; and by engineering the soil and litter layer. These processes can regulate the frequency, intensity and extent of fire. The evidence for these effects is strongest in environments with intermediate rainfall, warm temperatures and graminoid-dominated ground vegetation. Probably, extinction of Quaternary megafauna triggered increased biomass burning in many such environments. Recent and continuing declines of large vertebrates are likely to be significant contributors to changes in fire regimes and vegetation that are currently being experienced in many parts of the world. To date, rewilding projects that aim to restore large herbivores have paid little attention to the value of large animals in moderating fire regimes. Rewilding potentially offers a powerful tool for managing the risks of wildfire and its impacts on natural and human values.This article is part of the theme issue 'Trophic rewilding: consequences for ecosystems under global change'.

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.

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.