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.     

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.     

Herbaceous phytomass and nutrient concentrations of four grass species in Sudanian savanna woodland subjected to recurrent early fire

Fire is an integral ecological factor in African savanna ecosystems, but its effects on savanna productivity are highly variable and less understood. We conducted a field experiment to quantify changes in herbaceous phytomass and nutrient composition in a Sudanian savanna woodland subjected to annual early fire from 1993 to 2004. Fire effects were also assessed on two perennial and two annual grass species during the following growing season. Early fire significantly reduced above‐ground phytomass of the studied species (P = 0.03), their crude protein (P = 0.022), neutral detergent insoluble crude protein (P = 0.016) and concentrations of Ca, Fe and Mn (P < 0.05). Perennial grasses had higher above‐ground phytomass but lower total crude protein and fat than annual grasses. Nonstructural carbohydrates tended to be higher for annuals, while fibre and lignin contents were high for perennials. Except Na and Fe, the concentration of mineral elements varied between species. Fire did not affect measures of digestibility and metabolizable energy, but its effect differed significantly among species. In conclusion, the results illustrate that long‐term frequent fire will counterbalance the short‐term increase in soil fertility and plant nutrient concentrations claimed to be accrued from single or less frequent fire.     

When is a ‘forest’ a savanna,and why does it matter?

Savannas are defined based on vegetation structure, the central concept being a discontinuous tree cover in a continuous grass understorey. However, at the high‐rainfall end of the tropical savanna biome, where heavily wooded mesic savannas begin to structurally resemble forests, or where tropical forests are degraded such that they open out to structurally resemble savannas, vegetation structure alone may be inadequate to distinguish mesic savanna from forest. Additional knowledge of the functional differences between these ecosystems which contrast sharply in their evolutionary and ecological history is required. Specifically, we suggest that tropical mesic savannas are predominantly mixed tree–C₄ grass systems defined by fire tolerance and shade intolerance of their species, while forests, from which C₄ grasses are largely absent, have species that are mostly fire intolerant and shade tolerant. Using this framework, we identify a suite of morphological, physiological and life‐history traits that are likely to differ between tropical mesic savanna and forest species. We suggest that these traits can be used to distinguish between these ecosystems and thereby aid their appropriate management and conservation. We also suggest that many areas in South Asia classified as tropical dry forests, but characterized by fire‐resistant tree species in a C₄ grass‐dominated understorey, would be better classified as mesic savannas requiring fire and light to maintain the unique mix of species that characterize them.     

Vegetation and landscape connectivity control wildfire intervals in unmanaged semi‐arid shrublands and woodlands in Australia

Aim The aim of this study was to determine how spatial variation in vegetation type and landscape connectivity influence fire intervals in a semi‐arid landscape with low relief and complex mosaics of woodland and shrubland vegetation. Location Our study focused on a 15,500‐km² area of relatively undisturbed and unmanaged land in south‐western Australia, referred to as the Lake Johnston region. Methods We modelled fire‐interval data from a 67‐year (1940–2006) digital fire history database using a two‐parameter Weibull function, and tested for the effects of vegetation type and landscape connectivity on estimates of the length of fire intervals (Weibull parameter b) and the dependence of fire intervals on fuel age (Weibull parameter c). Results Vegetation type and landscape connectivity significantly influenced fire interval probability distributions. Fire intervals in shrublands (dense low shrub assemblage) were typically shorter (b = 46 years) and more dependent on fuel age (c = 2.33) than most other vegetation types, while fire intervals in open eucalypt woodlands were much longer (b = 405 years) and were less dependent on fuel age (c = 1.36) than in shrub‐dominated vegetation types. Areas adjacent to or surrounded by salt lakes burnt less frequently (b = 319 years) and fire intervals were less dependent on fuel age (c = 1.48) compared with more exposed areas (e.g. b < 101 years, c > 1.68). Fire intervals in thickets (dense tall shrub assemblage) were longer (b = 101 years) than would be expected from fuel loads, most likely because they were protected from fire by surrounding fuel‐limited woodlands. Main conclusions Fire intervals in south‐western Australia are strongly influenced by spatial variation in vegetation (fuel structure) and landscape connectivity. The importance of fuel structure as a control of fire intervals in south‐western Australia contrasts with other landscapes, where topographical gradients or climatic influences may override the effects of underlying vegetation. We found that, regardless of low relief, topographical features such as large salt lake systems limited the connectivity and spread of fire among landscape units in an analogous manner to lakes or mountainous features elsewhere.     

Fire regimes and phytomass growth dynamics in a Quercus coccifera garrigue

An experiment was set up in a Quercus coccifera garrigue in southern France to analyze the effect of burning frequency and season on phytomass production. Fire regimes consisted of late spring or early autumn burns, every 6 yr, every 3 yr, or every 2 yr. The experiment started in 1969 and lasted for 19 yr. In May 1981 and May 1987, 10 samples, each 1 m², were harvested per treatment. Fire frequency had an effect on the quantity of phytomass which was produced: above-ground phytomass decreased with increasing fire frequency. This was mainly due to the lower biomass of woody plants. In all burning treatments the phytomass of herbs was higher than in the unburned vegetation. Within each burning frequency, the total phytomass of the spring-burned vegetation was always higher than that of the autumn-burned community. Generally, the herb phytomass produced was higher in the autumn-burned plots. There were two fairly distinct phases in the period following fire, each with a different level of annual phytomass production. For the first six years it was about 300 g m^?2 yr^?1, falling thereafter to about 50 g m^?2 yr^?1. This and other studies on Q. coccifera garrigue indicate that this community is very resilient with respect to fire, but possesses a low productive capacity and does not show any sign of degeneration up to 30 yr old.     

Biomass consumption by surface fires across Earth's most fire prone continent

Landscape fire is a key but poorly understood component of the global carbon cycle. Predicting biomass consumption by fire at large spatial scales is essential to understanding carbon dynamics and hence how fire management can reduce greenhouse gas emissions and increase ecosystem carbon storage. An Australia‐wide field‐based survey (at 113 locations) across large‐scale macroecological gradients (climate, productivity and fire regimes) enabled estimation of how biomass combustion by surface fire directly affects continental‐scale carbon budgets. In terms of biomass consumption, we found clear trade‐offs between the frequency and severity of surface fires. In temperate southern Australia, characterised by less frequent and more severe fires, biomass consumed per fire was typically very high. In contrast, surface fires in the tropical savannas of northern Australia were very frequent but less severe, with much lower consumption of biomass per fire (about a quarter of that in the far south). When biomass consumption was expressed on an annual basis, biomass consumed was far greater in the tropical savannas (>20 times that of the far south). This trade‐off is also apparent in the ratio of annual carbon consumption to net primary production (NPP). Across Australia's naturally vegetated land area, annual carbon consumption by surface fire is equivalent to about 11% of NPP, with a sharp contrast between temperate southern Australia (6%) and tropical northern Australia (46%). Our results emphasise that fire management to reduce greenhouse gas emissions should focus on fire prone tropical savanna landscapes, where the vast bulk of biomass consumption occurs globally. In these landscapes, grass biomass is a key driver of frequency, intensity and combustion completeness of surface fires, and management actions that increase grass biomass are likely to lead to increases in greenhouse gas emissions from savanna fires.     

Self‐thinning and Tree Competition in Savannas

This paper examines the feasibility of applying self‐thinning concepts to savannas and how competition with herbaceous vegetation may modify self‐thinning patterns among woody plants in these ecosystems. Competition among woody plants has seldom been invoked as a major explanation for the persistence of herbaceous vegetation in mixed tree–grass ecosystems. On the contrary, the primary resource‐based explanations for tree–grass coexistence are based on tree–grass competition (niche‐separation) that assumes that trees are inferior competitors unless deeper rooting depths provide them exclusive access to water. Alternative nonresource‐based hypotheses postulate that trees are the better competitors, but that tree populations are suppressed by mortality related to fire, herbivores, and other disturbances. If self‐thinning of woody plants can be detected in savannas, stronger evidence for resource‐limitation and competitive interactions among woody plants would suggest that the primary models of savannas need to be adjusted. We present data from savanna sites in South Africa to suggest that self‐thinning among woody plants can be detected in low‐disturbance situations, while also showing signs that juvenile trees, more so than adults, are suppressed when growing with herbaceous vegetation in these ecosystems. This finding we suggest is evidence for size‐asymmetric competition in savannas.     

Influence of fire history on small mammal distributions: insights from a 100‐year post‐fire chronosequence

Aim Fire affects the structure and dynamics of ecosystems world‐wide, over long time periods (decades and centuries) and at large spatial scales (landscapes and regions). A pressing challenge for ecologists is to develop models that explain and predict faunal responses to fire at broad temporal and spatial scales. We used a 105‐year post‐fire chronosequence to investigate small mammal responses to fire across an extensive area of ‘tree mallee’ (i.e. vegetation characterized by small multi‐stemmed eucalypts). Location The Murray Mallee region (104,000 km²) of semi‐arid Australia. Methods First, we surveyed small mammals at 260 sites and explored the fire responses of four species using nonlinear regression models. Second, we assessed the predictive accuracy of models using cross‐validation and by testing with independent data. Third, we examined our results in relation to an influential model of animal succession, the habitat accommodation model. Results Two of four study species showed a clear response to fire history. The distribution of the Mallee Ningaui Ningaui yvonneae, a carnivorous marsupial, was strongly associated with mature vegetation characterized by its cover of hummock grass. The occurrence of breeding females was predicted to increase up to 40–105 years post‐fire, highlighting the extensive time periods over which small mammal populations may be affected by fire. Evaluation of models for N. yvonneae demonstrated that accurate predictions of species occurrence can be made from fire history and vegetation data, across large geographical areas. The introduced House Mouse Mus domesticus was the only species positively associated with recently burnt vegetation. Main conclusions Understanding the impact of fire over long time periods will benefit ecological and conservation management. In this example, tracts of long‐unburnt mallee vegetation were identified as important habitat for a fire‐sensitive native mammal. Small mammal responses to fire can be predicted accurately at broad spatial scales; however, a conceptual model of post‐fire change in community structure developed in temperate Australia is not, on its own, sufficient for small mammals in semi‐arid systems.     

Post‐fire habitat use of the golden‐backed tree‐rat (Mesembriomys macrurus) in the northwest Kimberley,Western Australia

Fire regimes are changing throughout the world. Changed fire patterns across northern Australian savannas have been proposed as a factor contributing to recent declines of small‐ and medium‐sized mammals. Despite this, few studies have examined the mechanisms that underpin how species use habitat in fire‐affected landscapes. We determined the habitats and resources important to the declining golden‐backed tree‐rat (Mesembriomys macrurus) in landscapes partially burnt by recent intense fire. We aimed to (i) compare the relative use of rainforest and savanna habitats; (ii) examine the effect of fire history on use of savanna habitats; and (iii) identify key foraging and denning resources. Habitat selection was examined by comparing the availability of eight habitat types around real (used) and generated (available) location points. Individuals used a range of habitats, but consistently selected long unburnt rainforest in preference to recently burnt savanna (1–12 months post‐fire); however, recently burnt savanna was used in preference to long unburnt savanna. Tree‐rats foraged in Terminalia hadleyana, Planchonia rupestris, Celtis philippensis and Owenia vernicosa, tree species that are found in a variety of habitat types. Individuals used a range of den sites, including cliffs, trees, logs, scree and stags found throughout the study area. Although multiple factors may have led to the decline of Mes. macrurus across its range, these results are consistent with the idea that changes in the savanna structure as a consequence of contemporary fire patterns could also have a role. The continued persistence of Mes. macrurus in the northwest Kimberley may be supported by land management strategies that conserve fruiting and hollow‐bearing trees, and maintain the availability of fire‐sensitive vegetation types.     

Does alpine grazing reduce blazing? A landscape test of a widely‐held hypothesis

Abstract ‘Alpine grazing reduces blazing’ is a widely and strongly held view concerning the effects of livestock grazing on fuels, and therefore fire behaviour and impact, in Australia's high country landscapes. As a test of this hypothesis, we examined the patterns of burning across the alpine (treeless) landscapes of the Bogong High Plains in Victoria, following the extensive fires of January 2003. Data were collected from multiple transects, each 3–5 km long, with survey points located randomly at either 50, 200 or 500 m intervals. The transects traversed the major regions of the Bogong High Plains, both grazed and ungrazed. At each point, we recorded whether the point was burnt or unburnt, the vegetation type (closed‐heath, open‐heath, grassland or herbfield), the estimated prefire shrub cover, slope, aspect, and a GPS location. At burnt heathland sites, we recorded the minimum twig diameter (an a posteriori measure of fire severity) in a sample of common shrubs. In total, there were 108 km of transect lines, 419 survey points and 4050 twig measurements, with sample points equally distributed across grazed and ungrazed country. The occurrence of fire (i.e. burnt or unburnt) in grazed and ungrazed areas was analysed by logistic regression; the variation in twig diameters by anova . Approximately half of all points were burnt. There was no statistically significant difference between grazed and ungrazed areas in the proportion of points burnt. Fire occurrence was determined primarily by vegetation type, with the proportion burnt being 0.87 for closed‐heath, 0.59 for open‐heath, and 0.13 for grassland and all snow‐patch herbfield points unburnt. In both closed‐heath and open‐heath, grazing did not significantly lower the severity of fire, as measured by the diameter of burnt twigs. We interpret the lack of a grazing effect in terms of shrub dynamics (little or no grazing effect on long‐term cover of taller shrubs), diet and behaviour of cattle (herbs and dwarf shrubs eaten; tall shrubs not eaten and closed‐heath vegetation generally avoided), and fuel flammability (shrubs more flammable than grass). Whatever effects livestock grazing may have on vegetation cover, and therefore fuels in alpine landscapes, they are likely to be highly localized, with such effects unlikely to translate into landscape‐scale reduction of fire occurrence or severity. The use of livestock grazing in Australian alpine environments as a fire abatement practice is not justified on scientific grounds.     

Long‐term variability and rainfall control of savanna fire regimes in equatorial East Africa

Fires burning the vast grasslands and savannas of Africa significantly influence the global carbon cycle. Projecting the impacts of future climate change on fire‐mediated biogeochemical processes in these dry tropical ecosystems requires understanding of how various climate factors influence regional fire regimes. To examine climate–vegetation–fire linkages in dry savanna, we conducted macroscopic and microscopic charcoal analysis on the sediments of the past 25 000 years from Lake Challa, a deep crater lake in equatorial East Africa. The charcoal‐inferred shifts in local and regional fire regimes were compared with previously published reconstructions of temperature, rainfall, seasonal drought severity, and vegetation dynamics to evaluate millennial‐scale drivers of fire occurrence. Our charcoal data indicate that fire in the dry lowland savanna of southeastern Kenya was not fuel‐limited during the Last Glacial Maximum (LGM) and Late Glacial, in contrast to many other regions throughout the world. Fire activity remained high at Lake Challa probably because the relatively high mean‐annual temperature (~22 °C) allowed productive C₄ grasses with high water‐use efficiency to dominate the landscape. From the LGM through the middle Holocene, the relative importance of savanna burning in the region varied primarily in response to changes in rainfall and dry‐season length, which were controlled by orbital insolation forcing of tropical monsoon dynamics. The fuel limitation that characterizes the region's fire regime today appears to have begun around 5000–6000 years ago, when warmer interglacial conditions coincided with prolonged seasonal drought. Thus, insolation‐driven variation in the amount and seasonality of rainfall during the past 25 000 years altered the immediate controls on fire occurrence in the grass‐dominated savannas of eastern equatorial Africa. These results show that climatic impacts on dry‐savanna burning are heterogeneous through time, with important implications for efforts to anticipate future shifts in fire‐mediated ecosystem processes.     

Micro-Scale Restoration: A 25-Year History of a Southern Illinois Barrens

We studied vegetation change of a remnant barrens in southern Illinois over twenty‐five years. The study area was periodically burned between 1969 and 1993, but fire was excluded for a 16‐year period (1974–1989). During the study, the barrens supported a mixture of species whose preferred habitats ranged from prairie and open woodlands to closed forest communities. The herbaceous vegetation may be on a trajectory characterized by increasing dominance of woodland species and declining prairie species. Fire management temporarily reversed this trend, but it continued once fire was excluded. Reintroduction of prescribed burning in 1990–1993 altered the vegetation trajectory but not back toward a species composition comparable to that present on the site before cessation of fire management after 1973. Following interruption of prescribed burning, tree basal area more than doubled, and density showed a 67% increase between premanagement conditions in 1968 and 1988. Salix humilis (prairie willow) density had significant negative correlations with tree density and basal area. However, there was no consistency in response of shrub species on the site to the varied site conditions over time. Fire management on the site may not recover the historic barrens that occurred on the site. Nevertheless, consistent fire management will drive vegetation changes toward increasing abundance of prairie and open woodland species that would otherwise be lost without burning.     

Tropical and north Pacific teleconnections influence fire regimes in pine-dominated forests of north-eastern California, USA

Aim To assess the importance of drought and teleconnections from the tropical and north Pacific Ocean on historical fire regimes and vegetation dynamics in north‐eastern California. Location The 700 km² study area was on the leeward slope of the southern Cascade Mountains in north‐eastern California. Open forests of ponderosa pine (Pinus ponderosa var. ponderosa Laws.) and Jeffrey pine (P. jeffreyi Grev. & Balf) surround a network of grass and shrub‐dominated meadows that range in elevation from 1650 to 1750 m. Methods Fire regime characteristics (return interval, season and extent) were determined from crossdated fire scars and were compared with tree‐ring based reconstructions of precipitation and temperature and teleconnections for the period 1700–1849. The effect of drought on fire regimes was determined using a tree‐ring based proxy of climate from five published chronologies. The number of forest‐meadow units that burned was compared with published reconstructions of the El Niño/Southern Oscillation (ENSO) and the Pacific Decadal Oscillation (PDO). Results Landscape scale fires burned every 7–49 years in meadow‐edge forests and were influenced by variation in drought, the PDO and ENSO. These widespread fires burned during years that were dryer and warmer than normal that followed wetter and cooler years. Less widespread fires were not associated with this wet, then dry climate pattern. Widespread fires occurred during El Niño years, but fire extent was mediated by the phase of the PDO. Fires were most widespread when the PDO was in a warm or normal phase. Fire return intervals, season and extent varied at decadal to multi‐decadal time scales. In particular, an anomalously cool, wet period during the early 1800s resulted in widespread fires that occurred earlier in the year than fires before or after. Main conclusions Fire regimes in north‐eastern California were strongly influenced by regional and hemispheric‐scale climate variation. Fire regimes responded to variation that occurred in both the north and tropical Pacific. Near normal modes of the PDO may influence fire regimes more than extreme conditions. The prevalence of widespread teleconnection‐driven fires in the historic record suggests that variation in the Pacific Ocean was a key regulator of fire regimes through its influence on local fuel production and successional dynamics in north‐eastern California.     

Seasonal leaf dynamics across a tree density gradient in a Brazilian savanna

Interactions between trees and grasses that influence leaf area index (LAI) have important consequences for savanna ecosystem processes through their controls on water, carbon, and energy fluxes as well as fire regimes. We measured LAI, of the groundlayer (herbaceous and woody plants <1-m tall) and shrub and tree layer (woody plants >1-m tall), in the Brazilian cerrado over a range of tree densities from open shrub savanna to closed woodland through the annual cycle. During the dry season, soil water potential was strongly and positively correlated with grass LAI, and less strongly with tree and shrub LAI. By the end of the dry season, LAI of grasses, groundlayer dicots and trees declined to 28, 60, and 68% of mean wet-season values, respectively. We compared the data to remotely sensed vegetation indices, finding that field measurements were more strongly correlated to the enhanced vegetation index (EVI, r ²=0.71) than to the normalized difference vegetation index (NDVI, r ²=0.49). Although the latter has been more widely used in quantifying leaf dynamics of tropical savannas, EVI appears better suited for this purpose. Our ground-based measurements demonstrate that groundlayer LAI declines with increasing tree density across sites, with savanna grasses being excluded at a tree LAI of approximately 3.3. LAI averaged 4.2 in nearby gallery (riparian) forest, so savanna grasses were absent, thereby greatly reducing fire risk and permitting survival of fire-sensitive forest tree species. Although edaphic conditions may partly explain the larger tree LAI of forests, relative to savanna, biological differences between savanna and forest tree species play an important role. Overall, forest tree species had 48% greater LAI than congeneric savanna trees under similar growing conditions. Savanna and forest species play distinct roles in the structure and dynamics of savanna–forest boundaries, contributing to the differences in fire regimes, microclimate, and nutrient cycling between savanna and forest ecosystems.     

Tree–grass coexistence in the Brazilian cerrado: demographic consequences of environmental instability

Aim Tropical savanna ecosystems are uniquely characterized by the co‐dominance of both trees and grasses. An operational understanding of the ecological processes involved in maintaining this condition is essential for understanding both the functioning of savanna systems as well as their potential response to environmental change. A simple model is presented to explore the potential for a demographic mechanism of long‐term tree persistence and temporal physiognomic stability in the Brazilian cerrado. Location The model is developed based on data from the humid cerrado of Brazil. Methods In contrast to many existing models of tree–grass dynamics a model is presented which is based on data from the humid cerrado of Brazil, which is both qualitatively and quantitatively different from many of the more arid savannas of the palaeotropics. The model focuses on the dynamics of a synthetic tree population, with particular attention given to reproduction, seedling establishment and fire effects; with separate sub‐models for grass production, fire and rainfall. Results The model successfully predicts coexistence across the full range of observed vegetation physiognomies, but only under limited conditions. Under coexistence conditions, the dynamics of the tree population are characterized by long periods of gradual decline, punctuated by occasional bursts of growth. However, in agreement with earlier studies, the model consistently over‐predicts domination by the tree component. Fire is identified as an overriding factor in determining model behaviour, and the response of reproduction and sapling recruitment to variance in the frequency of fire ignition is identified to be of potential importance in the functioning of the Brazilian cerrado. The key dynamics of the model which promote tree–grass coexistence are consistent with a number of established determinants of ecological resilience in savanna systems. Main conclusions The model identifies the importance of the effective exploitation of rare opportunities for favourable recruitment (e.g. exclusion from fire) by the tree population, in promoting coexistence within a predominantly adverse environment. Support is provided for an alternative demographic mechanism of tree–grass coexistence in the cerrado (the storage effect), which is not based on the limiting assumption of niche partitioning through differences in rooting depth. The results are consistent with those presented by recent modelling work based on the more arid savannas of southern Africa. The model presented here differs in the emphasis given to particular environmental and life‐history attributes which are critical in determining the tree–grass balance, but provides further general support for the potential role of demographic mechanisms (such as the storage effect) in determining the structure of tropical savannas. Despite having clear limitations, models can serve as valuable heuristic tools to aid the integration and exploration of existing data sets as well as our present understanding of key ecological processes.     

Herbivore–vegetation feedbacks can expand the range of savanna persistence: insights from a simple theoretical model

Theoretical models of tree–grass coexistence in savannas have focused primarily on the role of resource availability and fire. It is clear that herbivores heavily impact vegetation structure in many savannas, but their role in driving tree–grass coexistence and the stability of the savanna state has received less attention. Theoretical models of tree–grass dynamics tend to treat herbivory as a constant rather than a dynamic variable, yet herbivores respond dynamically to changes in vegetation structure in addition to modifying it. In particular, many savannas host two distinct herbivore guilds, grazers and browsers, both of which have the potential to exert profound effects on tree/grass balance. For example, grazers may indirectly favor tree recruitment by suppressing the destructive effects of fire, and browsers may facilitate the expansion of grassland by reducing the competitive dominance of trees. We use a simple theoretical model to explore the role of grazer and browser dynamics on savanna vegetation structure and stability across fire and resource availability gradients. Our model suggests that herbivores may expand the range of conditions under which trees and grasses are able to stably coexist, as well as having positive reciprocal effects on their own niche spaces. In addition, we suggest that given reasonable assumptions, indirect mutualisms can arise in savannas between functional groups of herbivores because of the interplay of consumption and ecosystem feedbacks.     

Variable rainfall has a greater effect than fire on the demography of the dominant tree in a semi‐arid Eucalyptus savanna

Rainfall, fire and competition are emphasized as determinants of the density and basal area of woody vegetation in savanna. The semi‐arid savannas of Australia have substantial multi‐year rainfall deficits and insufficient grass fuel to carry annual fire in contrast to the mesic savannas in more northern regions. This study investigates the influence of rainfall deficit and excess, fire and woody competition on the population dynamics of a dominant tree in a semi‐arid savanna. All individuals of Eucalyptus melanophloia were mapped and monitored in three, 1‐ha plots over an 8.5 year period encompassing wet and dry periods. The plots were unburnt, burnt once and burnt twice. A competition index incorporating the size and distance of neighbours to target individuals was determined. Supplementary studies examined seedling recruitment and the transition of juvenile trees into the sapling layer. Mortality of burnt seedlings was related to lignotuber area but the majority of seedlings are fire resistant within 12 months of germination. Most of the juveniles (≤1 cm dbh) of E. melanophloia either died in the dry period or persisted as juveniles throughout 8.5 years of monitoring. Mortality of juveniles was positively related to woody competition and was higher in the dry period than the wet period. The transition of juveniles to a larger size class occurred at extremely low rates, and a subsidiary study along a clearing boundary suggests release from woody competition allows transition into the sapling layer. From three fires the highest proportion of saplings (1–10 cm dbh) reduced to juveniles was only 5.6% suggesting rates of ‘top‐kill’ of E. melanophloia as a result of fire are relatively low. Girth growth was enhanced in wet years, particularly for larger trees (>10 cm dbh), but all trees regardless of size or woody competition levels are vulnerable to drought‐induced mortality. Overall the results suggest that variations in rainfall, especially drought‐induced mortality, have a much stronger influence on the tree demographics of E. melanophloia in a semi‐arid savanna of north‐eastern Australia than fire.     

Drivers of fire occurrence in a mountainous Brazilian cerrado savanna: Tracking long-term fire regimes using remote sensing

Fire is a natural disturbance in savannas, and defines vegetation physiognomy and structure, often influencing species diversity. Fire activity is determined by a wide range of factors, including long and short term climatic conditions, climate seasonality, wind speed and direction, topography, and fuel biomass. In Brazil, fire shapes the structure and composition of cerrado savannas, and the impact of fire on vegetation dynamics is well explored, but the drivers of variation in fire disturbance across landscapes and over time are still poorly understood. We reconstructed 31 years of fire occurrence history in the Serra do Cipó region, a highly-diverse cerrado landscape, located in the southern portion of the Espinhaço mountain range, state of Minas Gerais, Southeastern Brazil. We mapped burn scars using a time series of Landsat satellite images from 1984 to 2014. Our questions were 1) How does fire occurrence vary in time and space across the Serra do Cipó cerrado landscape? 2) Which climatic drivers may explain the spatial and inter-annual variation in fire occurrence on this landscape? 3) Is fire occurrence in this cerrado landscape moisture-limited or fuel-limited? We evaluated the inter-annual variation and distribution of burned areas, and used linear models to explain this variation in terms of rainfall amount (determinant of fuel load production), seasonal rainfall distribution (determinant of dry fuel availability), abnormality of precipitation (Standardized Precipitation Index – SPI), and vegetation type (Enhanced Vegetation Index – EVI). Contrary to our expectations, annual rainfall volume was weakly and negatively correlated with burned area, and the strongest predictor of burned area was drought during the ignition season. The length of the dry season and the distribution of rain along the season determined ignition probability, increasing fire occurrence during the driest periods. We conclude that the mountain cerrado vegetation at Serra do Cipó has a moisture-dependent fire regime, in contrast to the fuel-dependent fire regimes described for African savannas. These findings imply that savannas at different continents may have different recovery and resilience capabilities when subjected to changes in the fire regime, caused by direct anthropogenic activities or indirectly through climatic changes. The possible effects of these changes on cerrado landscapes are still unknown, and future studies should investigate if currently observed fire regimes have positive or negative impacts on vegetation diversity, recovery, resilience and phenology, thus helping managers to include fire management as conservation measure.     

Contrasting demographics of tropical savanna and temperate forest eucalypts provide insight into how savannas and forests function. A case study using Corymbia clarksoniana from north‐eastern Australia

Eucalypts (Eucalyptus spp. and Corymbia spp.) dominate many communities across Australia, including frequently burnt tropical savannas and temperate forests, which receive less frequent but more intense fires. Understanding the demographic characteristics that allow related trees to persist in tropical savannas and temperate forest ecosystems can provide insight into how savannas and forests function, including grass–tree coexistence. This study reviews differences in critical stages in the life cycle of savanna and temperate forest eucalypts, especially in relation to fire. It adds to the limited data on tropical eucalypts, by evaluating the effect of fire regimes on the population biology of Corymbia clarksoniana, a tree that dominates some tropical savannas of north‐eastern Australia. Corymbia clarksoniana displays similar demographic characteristics to other tropical savanna species, except that seedling emergence is enhanced when seed falls onto recently burnt ground during a high rainfall period. In contrast to many temperate forest eucalypts, tropical savanna eucalypts lack canopy‐stored seed banks; time annual seed fall to coincide with the onset of predictable wet season rain; have very rare seedling emergence events, including a lack of mass germination after each fire; possess an abundant sapling bank; and every tropical eucalypt species has the ability to maintain canopy structure by epicormically resprouting after all but the most intense fires. The combination of poor seedling recruitment strategies, coupled with characteristics allowing long‐term persistence of established plants, indicate tropical savanna eucalypts function through the persistence niche rather than the regeneration niche. The high rainfall‐promoted seedling emergence of C. clarksoniana and the reduction of seedling survival and sapling growth by fire, support the predictions that grass–tree coexistence in savannas is governed by rainfall limiting tree seedling recruitment and regular fires limiting the growth of juvenile trees to the canopy.