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Summary: Vegetation–fire feedbacks are important for determining the distribution of forest and savanna. To understand how vegetation structure controls these feedbacks, we quantified flammability across gradients of tree density from grassland to forest in the Brazilian Cerrado.We experimentally burned 102 plots, for which we measured vegetation structure, fuels, microclimate, ignition success and fire behavior.Tree density had strong negative effects on ignition success, rate of spread, fire‐line intensity and flame height. Declining grass biomass was the principal cause of this decline in flammability as tree density increased, but increasing fuel moisture contributed. Although the response of flammability to tree cover often is portrayed as an abrupt, largely invariant threshold, we found the response to be gradual, with considerable variability driven largely by temporal changes in atmospheric humidity. Even when accounting for humidity, flammability at intermediate tree densities cannot be predicted reliably.Fire spread in savanna–forest mosaics is not as deterministic as often assumed, but may appear so where vegetation boundaries are already sharp. Where transitions are diffuse, fire spread is difficult to predict, but should become increasingly predictable over multiple fire cycles, as boundaries are progressively sharpened until flammability appears to respond in a threshold‐like manner. [ABSTRACT FROM AUTHOR]

The ability of vegetation to ameliorate or exacerbate environmental extremes can generate feedbacks that mediate the distribution of biomes. It has been suggested that feedbacks between vegetation and frost damage may be important for maintaining savanna, particularly at the edge of the tropics.We quantified frost damage and air temperature across a network of 30 permanent plots distributed across tropical savanna–forest boundaries in Brazil during an uncommonly hard frost.Tree cover strongly buffered temperatures during frost events, such that forest sites were up to 5°C warmer than nearby sites occupied by open shrub savanna. Consequently, in forest, woody plants were not damaged, but in savanna, there was extensive dieback of trees and shrubs. Within savanna, frost had disproportionately large effects on small individuals, likely due to colder temperatures near the ground and the lower thermal mass of thin stems. Across species, frost tolerance was strongly correlated with latitudinal range limit, revealing the importance of minimum temperature as a species filter at the regional scale. Counterintuitively, savanna species, which are adapted to open habitats where frost is more likely, were more vulnerable to cold than were forest species and experienced more than twice the amount of leaf damage as co‐occurring forest species.Synthesis. Frost reinforces the effects of fire on vegetation structure by preferentially damaging trees in open environments and generating high loads of dead, flammable fuels. However, frost does not currently occur with sufficient frequency in the region to maintain open savanna by itself. Nevertheless, it occurs with sufficient frequency and severity that it likely acts as an environmental filter and evolutionary selective factor for some of the most frost‐sensitive species. Frost reinforces savanna‐forest boundaries in southern Brazil by damaging trees and creating flammable conditions where tree density is low, and by damaging savanna tree species to a greater extent than co‐occurring forest species. Frost is sufficiently frequent and severe to act as an environmental filter and evolutionary selective factor for some of the most cold‐sensitive tree species. [ABSTRACT FROM AUTHOR]

Fire regimes in savannas and forests are changing over much of the world. Anticipating the impact of these changes requires understanding how plants are adapted to fire. In this study, we test whether fire imposes a broad selective force on a key fire-tolerance trait, bark thickness, across 572 tree species distributed worldwide. We show that investment in thick bark is a pervasive adaptation in frequently burned areas across savannas and forests in both temperate and tropical regions where surface fires occur. Geographic variability in bark thickness is largely explained by annual burned area and precipitation seasonality. Combining environmental and species distribution data allowed us to assess vulnerability to future climate and fire conditions: tropical rainforests are especially vulnerable, whereas seasonal forests and savannas are more robust. The strong link between fire and bark thickness provides an avenue for assessing the vulnerability of tree communities to fire and demands inclusion in global models. [ABSTRACT FROM AUTHOR]

Numerous predictions indicate rising CO2 will accelerate the expansion of forests into savannas. Although encroaching forests can sequester carbon over the short term, increased fires and drought-fire interactions could offset carbon gains, which may be amplified by the shift toward forest plant communities more susceptible to fire-driven dieback. We quantify how bark thickness determines the ability of individual tree species to tolerate fire and subsequently determine the fire sensitivity of ecosystem carbon across 180 plots in savannas and forests throughout the 2.2-million km2 Cerrado region in Brazil. We find that not accounting for variation in bark thickness across tree species underestimated carbon losses in forests by ~50%, totaling 0.22 PgC across the Cerrado region. The lower bark thicknesses of plant species in forests decreased fire tolerance to such an extent that a third of carbon gains during forest encroachment may be at risk of dieback if burned. These results illustrate that consideration of trait-based differences in fire tolerance is critical for determining the climate-carbon-fire feedback in tropical savanna and forest biomes. [ABSTRACT FROM AUTHOR]

Fire shapes the distribution of savanna and forest through complex interactions involving climate, resources and species traits. Based on data from central Brazil, we propose that these interactions are governed by two critical thresholds. The fire-resistance threshold is reached when individual trees have accumulated sufficient bark to avoid stem death, whereas the fire-suppression threshold is reached when an ecosystem has sufficient canopy cover to suppress fire by excluding grasses. Surpassing either threshold is dependent upon long fire-free intervals, which are rare in mesic savanna. On high-resource sites, the thresholds are reached quickly, increasing the probability that savanna switches to forest, whereas low-resource sites are likely to remain as savanna even if fire is infrequent. Species traits influence both thresholds; saplings of savanna trees accumulate bark thickness more quickly than forest trees, and are more likely to become fire resistant during fire-free intervals. Forest trees accumulate leaf area more rapidly than savanna trees, thereby accelerating the transition to forest. Thus, multiple factors interact with fire to determine the distribution of savanna and forest by influencing the time needed to reach these thresholds. Future work should decipher multiple environmental controls over the rates of tree growth and canopy closure in savanna. [ABSTRACT FROM AUTHOR]

Upland tropical forests have expanded and contracted in response to past climates, but it is not clear whether similar dynamics were exhibited by gallery (riparian) forests within savanna biomes. Because such forests generally have access to ample water, their extent may be buffered against changing climates. We tested the long-term stability of gallery forest boundaries by characterizing the border between gallery forests and savannas and tracing the presence of gallery forest through isotopic analysis of organic carbon in the soil profile. We measured leaf area index, grass vs. shrub or tree coverage, the organic carbon, phosphorus, nitrogen and calcium concentrations in soils and the carbon isotope ratios of soil organic matter in two transitions spanning gallery forests and savanna in a Cerrado ecosystem. Gallery forests without grasses typically show a greater leaf area index in contrast to savannas, which show dense grass coverage. Soils of gallery forests have significantly greater concentrations of organic carbon, phosphorus, nitrogen and calcium than those of savannas. Soil organic carbon of savannas is significantly more enriched in 13C compared with that of gallery forests. This difference in enrichment is in part caused by the presence of C4 grasses in savanna ecosystem and its absence in gallery forests. Using the 13C abundance as a signature for savanna and gallery forest ecosystems in 1 m soil cores, we show that the borders of gallery forests have expanded into the savanna and that this process initiated at least 3000–4000bp based on 14C analysis. Gallery forests, however, may be still expanding as we found more recent transitions according to 14C activity measurements. We discuss the possible mechanisms of gallery forest expansion and the means by which nutrients required for the expansion of gallery forest might accumulate. [ABSTRACT FROM AUTHOR]

Exotic grasses are becoming increasingly abundant in Neotropical savannas, with Melinis minutiflora Beauv. being particularly invasive. To better understand the consequences for the native flora, we performed a field study to test the effect of this species on the establishment, survival and growth of seedlings of seven tree species native to the savannas and forests of the Cerrado region of Brazil. Seeds of the tree species were sown in 40 study plots, of which 20 were sites dominated by M. minutiflora, and 20 were dominated by native grasses. The exotic grass had no discernable effect on initial seedling emergence, as defined by the number of seedlings present at the end of the first growing season. Subsequent seedling survival in plots dominated by M. minutiflora was less than half that of plots dominated by native species. Consequently, at the end of the third growing season, invaded plots had only 44% as many seedlings as plots with native grasses. Above-ground grass biomass of invaded plots was more than twice that of uninvaded plots, while seedling survival was negatively correlated with grass biomass, suggesting that competition for light may explain the low seedling survival where M. minutiflora is dominant. Soils of invaded plots had higher mean Ca, Mg and Zn, but these variables did not account for the higher grass biomass or the lower seedling survival in invaded plots. The results indicate that this exotic grass is having substantial effects on the dynamics of the tree community, with likely consequences for ecosystem structure and function. [ABSTRACT FROM AUTHOR]

Exotic grasses are a serious threat to biodiversity in the cerrado savannas of central Brazil. Of particular concern is the possible role they may have in impeding tree regeneration at gallery (riverine) forest edges and increasing fire intensity, thereby driving gallery forest retreat. Here we quantify the effect of roads and distance from gallery forests on the abundance of the African grass Melinis minutiflora Beauv. and test for an effect of this species on woody plant regeneration and leaf area index. Melinis was present at approximately 70% of the sites near gallery forest edges, with its frequency declining sharply at greater distances from the edge. Melinis frequency was 2.8 times greater where roads were present nearby. Leaf area index (LAI) of the ground layer was 38% higher where Melinis was present than where it was absent. LAI was strongly correlated to fine fuel mass ( r2 = 0.80), indicating higher fuel loads where Melinis was present. The abundance of tree and shrub species in the ground layer was negatively related to LAI and to the presence of Melinis. The greater fuel accumulation and reduced tree regeneration caused by Melinis may cause a net reduction in forest area by increasing fire intensity at the gallery forest edge and slowing the rate of forest expansion. [ABSTRACT FROM AUTHOR]

Understanding the impact of fire on the demography of savanna trees and shrubs is necessary for understanding human impacts in tropical savannas. In a replicated experiment, we studied the impact of fire and vegetation cover on survival and growth of two subshrubs (Periandra mediterranea and Protium ovatum), two shrubs (Miconia albicans and Rourea induta) and three trees (Myrsine guianensis, Piptocarpha rotundifolia and Roupala montana) of the Brazilian cerrado savannas. Burning increased complete mortality (i.e. death of the individual) of five of the seven species, but primarily among individuals with stem diameters <4 mm. Stem mortality (i.e. topkill) was much more prevalent, primarily affecting individuals with stem diameter <32 mm, though all species experienced some topkill in even the largest size classes. Fires of higher intensity (flame length >2 m) caused greater mortality and topkill than fires of lower intensity (flame length <2 m). Pre-burn vegetation density had little effect on survival or resprout size, but did affect subsequent growth rates. Four species had greater growth rates in open sites, whereas only one species had greater growth rates in dense sites. For the three tree species and one shrub, resprouting individuals did not reach the minimum reproductive size within 1 year of burning, while the other shrub and the two subshrubs were able to reach reproductive size during this time, indicating that growth form largely determines the population response to frequent burning. [Copyright &y& Elsevier]

The savannas (cerrado) of south-central Brazil are currently subjected to frequent anthropogenic burning, causing widespread reduction in tree density. Increasing concentrations of atmospheric CO2 could reduce the impact of such frequent burning by increasing the availability of nonstructural carbohydrate, which is necessary for resprouting. We tested the hypotheses that elevated CO2 stimulates resprouting and accelerates replenishment of carbohydrate reserves. Using a factorial experiment, seedlings of a common Brazilian savanna tree, Keilmeyera coriacea, were grown at 350 ppm and 700 ppm CO2 and at two nutrient levels. To simulate burning, the plants were either clipped at 15 weeks or were left unclipped. Among unclipped plants, CO2 and nutrients both stimulated growth, with no significant interaction between nutrient and CO2 effects. Among clipped plants, both CO2 and nutrients stimulated resprouting. However, there was a strong interaction between CO2 and nutrient effects, with CO2 having a significant effect only in the presence of high nutrient availability. Under elevated CO2, carbohydrate reserves remained at higher levels following clipping. Root total nonstructural carbohydrate remained above 36% in all treatments, so carbohydrate reserves did not limit regrowth. These results indicate that under elevated CO2 this species may be better able to endure the high frequency of anthropogenic burning in the Brazilian savannas. [ABSTRACT FROM AUTHOR]

Fire controls tree cover in many savannas by suppressing saplings through repeated topkill and resprouting, causing a demographic bottleneck. Tree cover can increase dramatically if even a small fraction of saplings escape this fire trap, so modeling and management of savanna vegetation should account for occasional individuals that escape the fire trap because they are "better" (i.e., they grow faster than average) or because they are "lucky" (they experience an occasional longer‐than‐average interval without fire or a below‐average fire severity). We quantified variation in growth rates and topkill probability in Quercus laevis (turkey oak) in longleaf pine savanna to estimate the percentage of stems expected to escape the fire trap due to variability in (1) growth rate, (2) fire severity, and (3) fire interval. For trees growing at the mean rate and exposed to the mean fire severity and the mean fire interval, no saplings are expected to become adults under typical fire frequencies. Introducing variability in any of these factors, however, allows some individuals to escape the fire trap. A variable fire interval had the greatest influence, allowing 8% of stems to become adults within a century. In contrast, introducing variation in fire severity and growth rate should allow 2.8% and 0.3% of stems to become adults, respectively. Thus, most trees that escape the fire trap do so because of luck. By chance, they experience long fire‐free intervals and/or a low‐severity fire when they are not yet large enough to resist an average fire. Fewer stems escape the fire trap by being unusually fast‐growing individuals. It is important to quantify these sources of variation and their consequences to improve understanding, prediction, and management of vegetation dynamics of fire‐maintained savannas. Here we also present a new approach to quantifying variation in fire severity utilizing a latent‐variable model of logistic regression. [ABSTRACT FROM AUTHOR]

Highlights: [•] Tropical grassy biomes are extensive and critical to human livelihoods. [•] The definition of these grassy systems is inconsistent and misleading. [•] We discuss novel threats linked to problems with biome definition. [•] The assumption that ‘more trees are better’ does not hold for tropical grassy biomes. [Copyright &y& Elsevier]