Effect of fire regime on plant abundance in a tropical eucalypt savanna of north‐eastern Australia

Abstract Changes in plant abundance within a eucalypt savanna of north‐eastern Australia were studied using a manipulative fire experiment. Three fire regimes were compared between 1997 and 2001: (i) control, savanna burnt in the mid‐dry season (July) 1997 only; (ii) early burnt, savanna burnt in the mid‐dry season 1997 and early dry season (May) 1999; and (iii) late burnt, savanna burnt in the mid‐dry season 1997 and late dry season (October) 1999. Five annual surveys of permanent plots detected stability in the abundance of most species, irrespective of fire regime. However, a significant increase in the abundance of several subshrubs, ephemeral and twining perennial forbs, and grasses occurred in the first year after fire, particularly after late dry season fires. The abundance of these species declined toward prefire levels in the second year after fire. The dominant grass Heteropogon triticeus significantly declined in abundance with fire intervals of 4 years. The density of trees (>2 m tall) significantly increased in the absence of fire for 4 years, because of the growth of saplings; and the basal area of the dominant tree Corymbia clarksoniana significantly increased over the 5‐year study, irrespective of fire regime. Conservation management of these savannas will need to balance the role of regular fires in maintaining the diversity of herbaceous species with the requirement of fire intervals of at least 4‐years for allowing the growth of saplings >2 m in height. Whereas late dry season fires may cause some tree mortality, the use of occasional late fires may help maintain sustainable populations of many grasses and forbs.     

Monsoon rain forest seedling dynamics, northern Australia: contrasts with regeneration in eucalypt-dominated savannas

Aim To explore: (1) the relative influences of site conditions, especially moisture relations, on pathways and rates of monsoon rain forest seedling and sapling regeneration, especially of canopy dominants, in northern Australia; and (2) contrasts between regeneration syndromes of dominant woody taxa in savannas and monsoon rain forest. Location Four monsoon rain forest sites, representative of regional major habitat and vegetation types, in Kakadu National Park, northern Australia. Methods A decadal study involved: (1) initial assessment over 2.5 years to explore within‐year variability in seed rain, dormant seed banks and seedling (< 50 cm height) dynamics; and (2) thereafter, monitoring of seedling and sapling (50 cm height to 5 cm d.b.h.) dynamics undertaken annually in the late dry season. On the basis of observations from this and other studies, regeneration syndromes of dominant monsoon rain forest taxa are contrasted with comparable information for dominant woody savanna taxa, Eucalyptus and Corymbia especially. Results Key observations from the monsoon rain forest regeneration dynamics study component are that: (1) peak seed rain inputs of rain forest taxa were observed in the wet season at perennially moist sites, whereas inputs at seasonally dry sites extended into, or peaked in, the dry season; (2) dormant soil seed banks of woody rain forest taxa were dominated by pioneer taxa, especially figs; (3) longevity of dormant seed banks of woody monsoon rain forest taxa, including figs, was expended within 3 years; (4) seedling recruitment of monsoon rain forest woody taxa was derived mostly from wet season seed rain with limited inputs from soil seed banks; (5) at all sites rain forest seedling mortality occurred mostly in the dry season; (6) rain forest seedling and sapling densities were consistently greater at moist sites; (7) recruitment from clonal reproduction was negligible, even following unplanned low intensity fires. Main conclusions By comparison with dominant savanna eucalypts, dominant monsoon rain forest taxa recruit substantially greater stocks of seedlings, but exhibit slower aerial growth and development of resprouting capacity in early years, lack lignotubers in mesic species, and lack capacity for clonal reproduction. The reliance on sexual as opposed to vegetative reproduction places monsoon rain forest taxa at significant disadvantage, especially slower growing species on seasonally dry sites, given annual–biennial fires in many north Australian savannas.     

Are the eucalypt and non-eucalypt components of Australian tropical savannas independent?

Eucalypts (Eucalyptus and Corymbia spp.) dominate (>60%) the tree biomass of Australia's tropical savannas but account for only a fraction (28%) of the tree diversity. Because of their considerable biomass and adaptation to environmental stressors, such as fire, the eucalypts may drive tree dynamics in these savannas, possibly to the exclusion of non-eucalypts. We evaluated whether the eucalypt and non-eucalypt components in tropical savannas are dependent so that changes in one component are matched by opposite trends in the other. Using tree inventory data from 127 savanna sites across the rainfall and fire frequency gradients, we found that eucalypt and non-eucalypt basal area and species richness had a negative relationship. This relationship was maintained across the rainfall gradient, with rainfall having a positive effect on the basal area and species richness of both components, but with a greater effect in non-eucalypts. Fire frequency negatively affected basal area, but not species richness, although basal area and species richness of eucalypts and non-eucalypts did not differ in their response to fire. Rainfall appears to set the upper bounds to woody biomass in these mesic savannas, while fire maintains woody biomass below carrying capacity and facilitates coexistence of the components. The magnitude of the component responses, particularly for non-eucalypts, is determined by rainfall, but their dependence is likely due to their differential response to both rainfall and fire, but not to competition for resources. Thus, while eucalypts dominate biomass overall, at high rainfall sites non-eucalypt basal area and diversity are highest, especially where fire frequency is low.     

Buffering the savanna: fire regimes and disequilibrium ecology in West Africa

According to contemporary ecological theory, the mechanisms governing tree cover in savannas vary by precipitation level. In tropical areas with mesic rainfall levels, savannas are unstable systems in which disturbances, such as fire, determine the ratio of trees to grasses. Precipitation in these so-called “disturbance-driven savannas” is sufficient to support forest but frequent disturbances prevent transition to a closed canopy state. Building on a savanna buffering model we argue that a consistent fire regime is required to maintain savannas in mesic areas. We hypothesize that the spatiotemporal pattern of fires is highly regular and stable in these areas. Furthermore, because tree growth rates in savannas are a function of precipitation, we hypothesize that savannas with the highest rainfall levels will have the most consistent fire pattern and the most intense fires—thus the strongest buffering mechanisms. We analyzed the spatiotemporal pattern of burning over 11 years for a large subset of the West African savanna using a moderate resolution imaging spectroradiometer active fire product to document the fire regime for three savanna belts with different precipitation levels. We used LISA analysis to quantify the spatiotemporal patterns of fires, coefficient of variance to quantify differences in peak fire dates, and center or gravity pathways to characterize the spatiotemporal patterns of the fires for each area. Our analysis confirms that spatiotemporal regularity of the fire regime is greater for mesic areas that for areas where precipitation is lower and that areas with more precipitation have more regular fire regimes.     

Winners and losers: tropical forest tree seedling survival across a West African forest–savanna transition

Forest encroachment into savanna is occurring at an unprecedented rate across tropical Africa, leading to a loss of valuable savanna habitat. One of the first stages of forest encroachment is the establishment of tree seedlings at the forest–savanna transition. This study examines the demographic bottleneck in the seedlings of five species of tropical forest pioneer trees in a forest–savanna transition zone in West Africa. Five species of tropical pioneer forest tree seedlings were planted in savanna, mixed/transition, and forest vegetation types and grown for 12 months, during which time fire occurred in the area. We examined seedling survival rates, height, and stem diameter before and after fire; and seedling biomass and starch allocation patterns after fire. Seedling survival rates were significantly affected by fire, drought, and vegetation type. Seedlings that preferentially allocated more resources to increasing root and leaf starch (starch storage helps recovery from fire) survived better in savanna environments (frequently burnt), while seedlings that allocated more resources to growth and resource‐capture traits (height, the number of leaves, stem diameter, specific leaf area, specific root length, root‐to‐shoot ratio) survived better in mixed/transition and forest environments. Larger (taller with a greater stem diameter) seedlings survived burning better than smaller seedlings. However, larger seedlings survived better than smaller ones even in the absence of fire. Bombax buonopozense was the forest species that survived best in the savanna environment, likely as a result of increased access to light allowing greater investment in belowground starch storage capacity and therefore a greater ability to cope with fire. Synthesis: Forest pioneer tree species survived best through fire and drought in the savanna compared to the other two vegetation types. This was likely a result of the open‐canopied savanna providing greater access to light, thereby releasing seedlings from light limitation and enabling them to make and store more starch. Fire can be used as a management tool for controlling forest encroachment into savanna as it significantly affects seedling survival. However, if rainfall increases as a result of global change factors, encroachment may be more difficult to control as seedling survival ostensibly increases when the pressure of drought is lifted. We propose B. buonopozense as an indicator species for forest encroachment into savanna in West African forest–savanna transitions.     

Soil temperature and depth of legume germination during early and late dry season fires in a tropical eucalypt savanna of north‐eastern Australia

Abstract Temperatures that significantly increase seed germination of some tropical legumes (i.e. 80–100°C) were documented in the topsoil during the passage of early (May) and late (October) dry season fires in a tropical eucalypt savanna of north‐eastern Australia. Elevated temperatures penetrated at least 30 mm into the soil during the higher‐intensity, late dry season fires, but were only detected at 10 mm during the early dry season fires. The depth from which germination of two native legume forbs Galactia tenuiflora and Indigofera hirsuta occurred was positively related to the temperature elevation in the topsoil and was greater after late compared with early dry season fires. A broader range in germination depth, resulting in higher seedling densities, was recorded for I hirsuta after late dry season fires. These results suggest that seedling emergence of native leguminous forbs is likely to occur at a greater density after late rather than early dry season fires in tropical eucalypt savannas of north‐eastern Australia. Therefore, the season of burning, as a result of its relationship to fire intensity, can influence species composition through its effect on seed germination.     

Effects of seed burial and fire on seedling and sapling recruitment, survival and growth of African savanna woody plant species

Seed predation and seedling mortality can act as strong demographic “bottlenecks” to sapling recruitment in African savanna woodlands. Fire also limits tree recruitment from saplings by suppressing their growth. I conducted field experiments with 13 woody plant species to assess the effects of seed burial on seedling emergence rates and effects of fire on seedling and sapling survival and growth rates over a period of 8 years at a savanna plot in central Zambia, southern Africa. Seed removal rates by small rodents varied among years and buried seeds had significantly higher emergence rates than seeds exposed to predators in most but not all the species. Annual burning reduced sapling growth in some species but in other species saplings experienced successive shoot die back even in the absence of fire. The findings show that for some woody species, seed predation is an important constraint to seedling recruitment but not for others and annual fires are important hindrances to demography and growth for some species but not others. Thus, demographic “bottlenecks” occur at different life history stages in different savanna woody species and these have the potential to alter woody tree competitive relationships and ultimately savanna structure.     

Fire resilience of ant assemblages in long‐unburnt savanna of northern Australia

Abstract Tropical savannas and rainforests contrast in their flammability and the fire resilience of their associated species. While savanna species generally exhibit high resilience to burning, there is much debate about the fire resilience of forest‐associated species, and the persistence of forest patches in a flammable savanna matrix. Where fire has been excluded, savanna tends on a trajectory towards forest, with an increase in forest‐associated plants and animal species. This study tested the idea that given the high proportion of forest‐associated taxa in long‐unburnt savanna, the fauna of these areas would be expected to exhibit less resilience to fire than the fauna in frequently burnt savannas. The study investigated the immediate and short‐term effects on ant assemblages of re‐introducing fire into long‐unburnt savanna in northern Australia. The ant fauna exhibited high resistance to fires, with no significant short‐term change in mean abundance or species richness; instead, seasonality had a far stronger influence on overall ant activity. Fire caused dramatic declines in dominance of the patchily distributed forest‐associated species Oecophylla smaragdina and Papyrius sp., but had no effect on overall dominance by open savanna species of Iridomyrmex. Dominance by Iridomyrmex pallidus declined, but this was compensated for by increases in I. reburrus, while two other species of Iridomyrmex showed no change. This indicates a high level of functional redundancy among dominant species of Iridomyrmex, which universally dominate open savanna communities, but not of dominant forest‐associated species. Overall, our findings demonstrate a high degree of fire‐resilience of the long‐unburnt savanna ant fauna. Despite the occurrence of forest‐associated species, the high proportion of savanna species persisting in this habitat means that long‐unburnt savanna retains the general response characteristics of frequently burnt savanna.     

Simply the best: the transition of savanna saplings to trees

Tree cover in savannas is determined as much by disturbances from fire and herbivory as by rainfall and soil resources. Fire especially acts to limit tree cover via a demographic bottleneck, limiting the recruitment of tree saplings to adults. Because sapling growth rates determine rates of sapling to tree recruitment, predicting changes in tree cover requires data on sapling growth rates, commonly expressed as population means. Here, we discuss the variability in sapling growth rates in Acacia populations in a savanna in Hluhluwe iMfolozi Park in South Africa. Saplings growing at mean rates under typical fire regimes in African savannas would likely never escape the fire‐trap to become adults. Only the fastest growing saplings could grow above the flame zone between fires. We suggest that maximum growth rates are more ecologically relevant than mean growth rates in natural populations and experiments. Maximum growth rates are better than mean growth rates as predictors of sapling release within species, as shown here, and probably of which species are likely ‘winners’ in savanna tree communities.     

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.     

Effects of Fire on Seedling Diversity and Plant Reproduction (Sexual vs. Vegetative) in Neotropical Savannas Differing in Tree Density

Little is known about the effects of fire on the structure and species composition of Neotropical savanna seedling communities. Such effects are critical for predicting long‐term changes in plant distribution patterns in these ecosystems. We quantified richness and density of seedlings within 144 plots of 1 m² located along a topographic gradient in long‐unburned (fire protected since 1983) and recently burned (September 2005) savannas in Brazil. These savannas differ in tree density and canopy cover. Sites along the gradient, however, did not differ in species composition prior to the fire. In recently burned savannas we also evaluated the contribution of vegetative reproduction relative to sexual reproduction by quantifying richness and density of root suckers. Finally, we tested seed tolerance to pulses of high temperatures—similar to those occurring during fires on the soil surface and below—of five dominant savanna tree species. Seedlings were more abundant and diverse in unburned than in burned savannas. Seedling species composition differed among unburned and burned savannas probably reflecting early differences in root: shoot biomass allocation patterns. In recently burned savannas, root suckers were more abundant and diverse than seedlings. Relatively long exposures (>10 min) of temperatures of 90 °C reduced seed germination in all studied species suggesting a negative effect of fire on germination of seeds located at or aboveground level. Because vegetative reproduction contributes more than sexual reproduction in burned environments, frequent fires are likely to cause major shifts in species composition of Neotropical savanna plant communities, favoring clonally produced recruits along tree density/topographic gradients.     

Does rapid utilization of elevated nutrient availability allow eucalypts to dominate in the tropical savannas of Australia?

Northern Australia's savannas are among the most fire‐prone biomes on Earth and are dominated by eucalypts (Eucalyptus and Corymbia spp.). It is not clear what processes allow this group to dominate under such extreme fire frequencies and whether a superior ability to compete for nutrients and water might play a role. There is evidence that eucalypts are adapted to frequent fires; juvenile eucalypts escape the fire trap by growing rapidly in height between fires. However, non‐eucalypts are less able to escape the fire trap and tend to have stand structures strongly skewed toward suppressed juveniles. The mechanisms that drive these contrasting fire responses are not well understood. Here, we describe the results of a controlled glasshouse seedling experiment that evaluated the relative importance of nutrient and water availability in determining height growth and biomass growth of two eucalypt and one noneucalypt tree species, common in northern Australian savannas. We demonstrate that growth of eucalypt seedlings is particularly responsive to nutrient addition. Eucalypt seedlings are able to rapidly utilize soil nutrients and accumulate biomass at a much greater rate than noneucalypt seedlings. We suggest that a seasonal spike in nutrient availability creates a nutrient‐rich microsite that allows eucalypt seedlings to rapidly gain height and biomass, increasing their likelihood of establishing successfully and reaching a fire‐resistant size. Our results extend our understanding of how eucalypts dominate northern Australian savannas under extremely high fire frequencies.     

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.     

The impact of experimental fire regimes on seed production in two tropical eucalypt species in northern Australia

Abstract This study investigated the effect of three experimental fire regimes on the fecundity, ovule development and seedfall of two common wet-dry tropical savanna eucalypts, Eucalyptus minima and Eucalyptus tetrodonta, in northern Australia. Both species flower early in the dry season and ovule development occurs during the dry season. This coincides with a period of frequent fires. The three fire regimes considered were applied for four years between 1990 and 1994. These regimes were (i) Unburnt, (ii) Early, fires lit early in the dry season, and (iii) Late, fires lit late in the dry season. The treatments were applied to nine catchments (15–20 km²) with each fire regime replicated three times. Fire intensity typically increases as the dry season proceeds. Therefore, early dry season fires generally differ from late dry season fires in both their intensity and their timing in relation to the reproductive phenology of the eucalypts. Late dry season burning significantly reduced the fecundity of both species, whereas Early burning had no significant effect. Ovule success was significantly reduced by the Early burning for both species. The Late burning significantly reduced ovule success in E. tetrodonta, but not in E. miniata. The results suggest that fire intensity and fire timing may both be important determinants of seed supply. Fire intensity may be a determinant of fecundity, whereas fire timing in relation to the reproduction phenology may have a significant impact on ovule survival. Both fire regimes resulted in a substantial reduction in seed supply compared with the Unburnt treatment. This may have a significant impact on seedling regeneration of these tropical savanna eucalypts.     

Bark thickness determines fire resistance of selected tree species from fire-prone tropical savanna in north Australia

We investigated the fire resistance conferred by bark of seven common tree species in north Australian tropical savannas. We estimated bark thermal conductance and examined the relative importance of bark thickness, density and moisture content for protecting the cambium from lethal fire temperatures. Eucalypt and non-eucalypt species were contrasted, including the fire-sensitive conifer Callitris intratropica. Cambial temperature responses to bark surface heating were measured using a modified wick-fire technique, which simulated a heat pulse comparable to surface fires of moderate intensity. Bark thickness was a better predictor of resistance to cambial injury from fires than either bark moisture or density, accounting for 68% of the deviance in maximum temperature of the cambium. The duration of heating required to kill the cambium of a tree (τ_c) was directly proportional to bark thickness squared. Although species did not differ significantly in their bark thermal conductance (k), the thinner barked eucalypts nevertheless achieved similar or only slightly lower levels of fire resistance than much thicker barked non-eucalypts. Bark thickness alone cannot account for the latter and we suggest that lower bark moisture content among the eucalypts also contributes to their apparent fire resistance. Unique eucalypt meristem anatomy and epicormic structures, combined with their bark traits, probably facilitate resprouting after fire and ensure the dominance of eucalypts in fire-prone savannas. This study emphasises the need to take into account both the thermal properties of bark and the mechanism of bud protection in characterising the resprouting ability of savanna trees.     

Shifts in functional traits elevate risk of fire‐driven tree dieback in tropical savanna and forest biomes

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.     

Decadal dynamics of tree cover in an Australian tropical savanna

Spatio‐temporal variation in tropical savanna tree cover remains poorly understood. We aimed to quantify the drivers of tree cover in tropical mesic savannas in Kakadu National Park by relating changes in tree cover over 40 years to: mean annual rainfall, fire activity, initial tree cover and prior changes in tree cover. Aerial photography, acquired in 1964, 1984 and 2004, was obtained for fifty sites in Kakadu that spanned a rainfall gradient from approximately 1200 to 1600 mm. The remotely sensed estimates of tree cover were validated via field survey. Linear mixed effects modelling and multi‐model inference were used to assess the strength and form of the relationships between tree cover and predictor variables. Over the 40 years, tree cover across these savannas increased on average by 4.94 ± 0.88%, but was spatio‐temporally variable. Tree cover showed a positive albeit weak trend across the rainfall gradient. The strength of this positive relationship varied over the three measurement times, and this suggests that other factors are important in controlling tree cover. Tree cover was positively related to prior tree cover, and negatively correlated with fire activity. Over 20 years tree cover was more likely to increase if (i) tree cover was initially low or (ii) had decreased in the previous 20‐year interval or (iii) there had been fewer fires. Across the examined rainfall gradient, the greater variability in fire activity and inherently higher average tree cover at the wetter latitudes resulted in greater dynamism of tree cover compared with the drier latitudes. This is consistent with savanna tree cover being determined by interactions between mean annual rainfall, tree competition and frequent fire in these tropical mesic savannas.     

Effects of Canopy Cover and Ground Disturbance on Establishment of an Invasive Grass in an Australia Savanna1

Andropogon gayanus (gamba grass) is an introduced pasture grass that threatens Australia's tropical savannas by modifying fire regimes and species composition. To understand the establishment requirements of A. gayanus, we undertook a field experiment to determine the effect of canopy cover and ground layer disturbance on seedling emergence and survival. Seed was sown under three canopy treatments (undisturbed, artificial canopy gap, and natural canopy gap) and under three ground layer treatments (Control, Vegetation disturbed, and Soil disturbed). Results have shown that A. gayanus can establish and survive regardless of canopy cover or ground disturbance, although such site disturbances will increase establishment success. Disturbance of both the overstorey canopy and the ground layer increased A. gayanus emergence, whereas seedling survival to 12 mo after seed sowing was affected by ground layer disturbance alone. Disturbance of the canopy increased light transmission, which may have promoted germination. Ground layer disturbance may also have increased light transmission and suitable sites for establishment, and reduced competition for resources, such as water and nutrients. The ability of A. gayanus to spread along disturbed areas, establish in relatively undisturbed savannas, and resprout after fire within 6 mo after seedling emergence suggests that this species will become increasingly widespread in Australia's tropical savannas. Its control is urgently required.     

Can savannas become forests? A coupled analysis of nutrient stocks and fire thresholds in central Brazil

Aims

The effects of fire ensure that large areas of the seasonal tropics are maintained as savannas. The advance of forests into these areas depends on shifts in species composition and the presence of sufficient nutrients. Predicting such transitions, however, is difficult due to a poor understanding of the nutrient stocks required for different combinations of species to resist and suppress fires.

Methods

We compare the amounts of nutrients required by congeneric savanna and forest trees to reach two thresholds of establishment and maintenance: that of fire resistance, after which individual trees are large enough to survive fires, and that of fire suppression, after which the collective tree canopy is dense enough to minimize understory growth, thereby arresting the spread of fire. We further calculate the arboreal and soil nutrient stocks of savannas, to determine if these are sufficient to support the expansion of forests following initial establishment.

Results

Forest species require a larger nutrient supply to resist fires than savanna species, which are better able to reach a fire-resistant size under nutrient limitation. However, forest species require a lower nutrient supply to attain closed canopies and suppress fires; therefore, the ingression of forest trees into savannas facilitates the transition to forest. Savannas have sufficient N, K, and Mg, but require additional P and Ca to build high-biomass forests and allow full forest expansion following establishment.

Conclusions

Tradeoffs between nutrient requirements and adaptations to fire reinforce savanna and forest as alternate stable states, explaining the long-term persistence of vegetation mosaics in the seasonal tropics. Low-fertility limits the advance of forests into savannas, but the ingression of forest species favors the formation of non-flammable states, increasing fertility and promoting forest expansion.     

Fire‐sensitive species dominate seed rain after fire suppression: Implications for plant community diversity and woody encroachment in the Cerrado

Woody encroachment is becoming common in tropical savannas. We studied natural seed rain and performed seed addition experiments in a Brazilian savanna that had not been burned for several decades. We found greater abundance of fire‐sensitive species in the seed rain, likely contributing to woody encroachment. Flexible fire management policies that allow for natural and prescribed fires may be required to maintain savanna diversity.