Grass feedbacks on fire stabilize savannas

Savannas commonly consist of a discontinuous cover of overstory trees and a groundcover of grasses. Savanna models have previously demonstrated that vegetation feedbacks on fire frequency can limit the density of overstory trees, thereby maintaining savannas. Positive feedbacks of either savanna trees alone or trees and grasses together on fire frequency have been shown to result in a stable savanna equilibrium. Grass feedbacks on fire frequency, in contrast, have resulted in stable equilibria in either a grassland or forest state, but not in a savanna. These results, however, were derived from a system of differential equations that assumes that fire occurrence is strictly deterministic and that vegetation losses due to fire are continuous in time. We develop an alternative formulation of the grass-fire feedback model that assumes that fires are discrete and occur stochastically in time to examine the influence of these assumptions on the predicted state of the system. We show that incorporating fire as a discrete event can produce a recurring temporal refuge in which both grass and trees co-occur in a stable, bounded savanna. In our model, tree abundance is limited without invoking demographic bottlenecks in the transition from fire-sensitive to fire-resistant life history stages. An increasing strength of grass feedback on fire results in regular, predictable fires, which suggests that the system can also be modeled using a set of difference equations. We implement this discrete system using modified Leslie/Gower difference equations and demonstrate the existence of a bounded savanna state in this model framework. Our results confirm the potential for grass feedbacks to result in stable savannas, and indicate the importance of modeling fire as a discrete event rather than as a loss rate that is continuous in time.     

Are savannas patch-dynamic systems? A landscape model

Savannas are ecosystems characterized by the coexistence of woody species (trees and bushes) and grasses. Given that savanna characteristics are mainly formed from competition, herbivory, fire, woodcutting, and patchy soil and precipitation characteristics, we propose a spatially explicit model to examine the effects of the above-mentioned parameters on savanna vegetation dynamics in space and time. Furthermore, we investigate the effects of the above-mentioned parameters on tree–bush–grass ratios, as well as the degrees of aggregation of tree–bush–grass biomass. We parameterized our model for an arid savanna with shallow soil depth as well as a mesic one with generally deeper and more variable soil depths. Our model was able to reproduce savanna vegetation characteristics for periods of time over 2000 years with daily updated time steps. According to our results, tree biomass was higher than bush biomass in the arid savanna but bush biomass exceeded tree and grass biomass in the simulated mesic savanna. Woody biomass increased in our simulations when the soil's porosity values were increased (mesic savanna), in combination with higher precipitation. Savanna vegetation varied from open savanna to woodland and back to open savanna again. Vegetation cycles varied over ∼300-year cycles in the arid and ∼220-year cycles in the mesic-simulated savanna. Autocorrelation values indicated that there are both temporal and spatial vegetation cycles. Our model indicated cycling savanna vegetation at the landscape scale, cycles in cells, and patchiness, i.e. patch dynamics.     

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

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

Chemical composition of rivers and streams in the forest and savanna zones of Nigeria

The aim of this study is to investigate the differences in the chemical conditions of lotic waterbodies in the two major ecosystems in Nigeria, the forest and savanna zones. The forest waters were slightly acidic (mean±SD pH = 6.72±0.58) while the savanna waters were slightly alkaline (pH = 7.11±0.33). The cationic order of dominance in the forest waters was Na⁺ > Ca²⁺ > Mg²⁺ > K⁺ in contrast to Ca²⁺ > Mg²⁺ > Na⁺ > K⁺ in savanna waters. The forest waters were chlorided (typical of coastal and/or marine waters) whereas the savanna waters were carbonated in nature, typical of the worldwide freshwater. Organic carbon was significantly higher in forest waters than in the savanna waters (p < 0.05) while nutrient compounds were significantly higher in savanna waters than in forest waters. The seasonal variation of the chemical parameters was generally more evident in savanna than in forest waters. The differences in water quality between the two major vegetation zones reflect the differences in the biogeochemical processes and nutrient cycling that characterise forest and savanna ecosystems.     

Simulating wildfire patterns using a small-world network model

This paper presents the development and validation results of a weighted small-world network model designed to simulate fire patterns in real heterogeneous landscapes. Fire spread is simulated on a gridded landscape, a mosaic in which each cell represents an area of the land surface. In this model, the interaction between burning and non-burning cells (here, due to flame radiation) may extend well beyond nearest neighbors, and depends on local conditions of wind, topography, and vegetation. An approach based on the coupling of the solid flame model with the Monte Carlo method is used to predict the radiative heat flux from the flame generated by the burning of each combustible cell to its neighbors. The weighting procedure takes into account latency (a combustible cell will only ignite when it has accumulated enough energy along time) and flaming persistence of burning cells. The model is applied to very different fire scenarios: a historical Mediterranean fire that occurred in southeastern France in 2005 and experimental fires conducted in arid savanna fuels in South Africa in 1992. Model results are found to be in agreement with real fire patterns, in terms both of rate of spread, and of the area and shape of the burn. This work also shows that the fractal properties of fire patterns predicted by the model are similar to those observed from satellite images of three other Mediterranean fire scars.     

N biogeochemistry and cycling in two well-drained savannas: a comparison between the Orinoco Basin (Llanos,Venezuela) and Ivory Coast (western Africa)

When a savanna burns, a decline in the input of organic matter and nutrients to the soil occurs. However, the existence of recurrent fires is a natural condition and N depletion by fire is not incompatible with the existence of savannas per se. Consequently, savanna vegetations have evolved under fires, implying a near to steady-state N budget. In some Australian and African ecosystems, N fixation appears to be insufficient to replace losses inducing soil-N depletion, whereas in neotropical Llanos and western African savannas, the N-fixation and precipitation seem to be enough to maintain production despite fires. This review presents information about well-drained savannas, namely Trachypogon savannas in Orinoco Llanos and Andropogonae savannas in Ivory Coast. The sites present similarities in climate and fire regime differing in soil parent material. A report on N budgets is presented. The budget was positive, since losses seem to be balanced by inputs in rainfall and biological fixation. Uncertainties in flux measures indicate that more work on those aspects are required. Results indicate that the estimated amounts of dinitrogen fixation and deposition seem to be enough to maintain plant production. Data support the hypothesis that the vegetation in the natural savanna has evolved under fire constraint.     

Effects of four decades of fire manipulation on woody vegetation structure in Savanna

The amount of carbon stored in savannas represents a significant uncertainty in global carbon budgets, primarily because fire causes actual biomass to differ from potential biomass. We analyzed the structural response of woody plants to long-term experimental burning in savannas. The experiment uses a randomized block design to examine fire exclusion and the season and frequency of burn in 192 7-ha experimental plots located in four different savanna ecosystems. Although previous studies would lead us to expect tree density to respond to the fire regime, our results, obtained from four different savanna ecosystems, suggest that the density of woody individuals was unresponsive to fire. The relative dominance of small trees was, however, highly responsive to fire regime. The observed shift in the structure of tree populations has potentially large impacts on the carbon balance. However, the response of tree biomass to fire of the different savannas studied were different, making it difficult to generalize about the extent to which fire can be used to manipulate carbon sequestration in savannas. This study provides evidence that savannas are demographically resilient to fire, but structurally responsive.     

Forest Restoration and Fire: Principles in the Context of Place

Abstract:  There is broad consensus that active management through thinning and fire is urgently needed in many forests of the western United States. This consensus stems from physically based models of fire behavior and substantial empirical evidence. But the types of thinning and fire and where they are applied are the subjects of much debate. We propose that low thinning is the most appropriate type of thinning practice. Treating surface fuels, reducing ladder fuels, and opening overstory canopies generally produce fire-safe forest conditions, but large, fire-resistant trees are also important components of fire-safe forests. The context of place is critical in assigning priority for the limited resources that will be available for restoration treatments. Historical low-severity fire regimes, because of their current high hazards and dominance by fire-resistant species, are the highest priority for treatment. Mixed-severity fire regimes are of intermediate priority, and high-severity fire regimes are of lowest priority. Classification systems based on potential vegetation will help identify these fire regimes at a local scale.     

Biotic and abiotic regulation of lightning fire initiation in the mixedwood boreal forest

Lightning fire is the dominant natural disturbance of the western mixedwood boreal forest of North America. We quantified the independent effects of weather and forest composition on lightning fire initiation (a detected and recorded fire start) patterns in Alberta, Canada, to demonstrate how these biotic and abiotic components contribute to ecosystem dynamics in the mixedwood boreal forest. We used logistic regression to describe variation in annual initiation occurrence among 10,000-ha landscape units (voxels) covering a 9 million-ha study region over 11 years. At a voxel scale, forest composition explained more variation in annual initiation than did weather indices. Initiations occurred more frequently in landscapes with more conifer fuels (Picea spp.), and less in aspen-dominated (Populus spp.) ones. Initiations were less frequent in landscapes that had recently burned. Variation in initiation was also influenced by joint weather-lightning indices, but to a lesser degree. For each voxel, these indices quantified the number of days in the fire season when moisture levels were low and lightning was detected. Regional indices of fire weather severity explained substantial interannual variation of initiation, and the effect of forest composition was stronger in years with more severe fire weather. Our study is a conclusive demonstration of biotic and abiotic regulation of lightning fire initiation in the mixedwood boreal forest. The independent effects of forest composition emphasize that vegetation feedbacks strongly regulate disturbance dynamics in the region.     

Modelling forest–savanna mosaic dynamics in man-influenced environments: effects of fire,climate and soil heterogeneity

Forests and savannas are the major ecotypes in humid tropical regions. Under present climatic conditions, forest is in a phase of natural expansion over savanna, but traditional human activities, especially fires, have strongly influenced the succession. We here present a new model, FORSAT, dedicated to the forest–savanna mosaic on a landscape scale and based on stochastic modelling of key processes (fire and succession cycle) and consistent with common field data. The model is validated by comparison between the qualitative emergent behaviour of the model and results of biogeographical field studies. Three types of forest succession are shown: progression of the forest edge, formation and coalescence of clumps in savanna and global afforestation of savanna. The parameters (frequency of savanna fires, climate and soil fertility) appear to have comparable effects and there is a sharp threshold between a forest edge progression scenario and the cluster formation one. Moreover, pioneer seed dispersal pattern and recruitment are determinant: peaked curves near a seed source and far dispersal combine to increase the fitness of the pioneers.     

Soil feedback of exotic savanna grass relates to pathogen absence and mycorrhizal selectivity

Enemy release of exotic plants from soil pathogens has been tested by examining plant-soil feedback effects in repetitive growth cycles. However, positive soil feedback may also be due to enhanced benefit from the local arbuscular mycorrhizal fungi (AMF). Few studies actually have tested pathogen effects, and none of them did so in arid savannas. In the Kalahari savanna in Botswana, we compared the soil feedback of the exotic grass Cenchrus biflorus with that of two dominant native grasses, Eragrostis lehmanniana and Aristida meridionalis. The exotic grass had neutral to positive soil feedback, whereas both native grasses showed neutral to negative feedback effects. Isolation and testing of root-inhabiting fungi of E. lehmanniana yielded two host-specific pathogens that did not influence the exotic C. biflorus or the other native grass, A. meridionalis. None of the grasses was affected by the fungi that were isolated from the roots of the exotic C. biflorus. We isolated and compared the AMF community of the native and exotic grasses by polymerase chain reaction-denaturing gradient gel elecrophoresis (PCR-DGGE), targeting AMF 18S rRNA. We used roots from monospecific field stands and from plants grown in pots with mixtures of soils from the monospecific field stands. Three-quarters of the root samples of the exotic grass had two nearly identical sequences, showing 99% similarity with Glomus versiforme. The two native grasses were also associated with distinct bands, but each of these bands occurred in only a fraction of the root samples. The native grasses contained a higher diversity of AMF bands than the exotic grass. Canonical correspondence analyses of the AMF band patterns revealed almost as much difference between the native and exotic grasses as between the native grasses. In conclusion, our results support the hypothesis that release from soil-borne enemies may facilitate local abundance of exotic plants, and we provide the first evidence that these processes may occur in arid savanna ecosystems. Pathogenicity tests implicated the involvement of soil pathogens in the soil feedback responses, and further studies should reveal the functional consequences of the observed high infection with a low diversity of AMF in the roots of exotic plants.     

Simulating dynamic and mixed-severity fire regimes: A process-based fire extension for LANDIS-II

Fire regimes result from reciprocal interactions between vegetation and fire that may be further affected by other disturbances, including climate, landform, and terrain. In this paper, we describe fire and fuel extensions for the forest landscape simulation model, LANDIS-II, that allow dynamic interactions among fire, vegetation, climate, and landscape structure, and incorporate realistic fire characteristics (shapes, distributions, and effects) that can vary within and between fire events. We demonstrate the capabilities of the new extensions using two case study examples with very different ecosystem characteristics: a boreal forest system from central Labrador, Canada, and a mixed conifer system from the Sierra Nevada Mountains (California, USA). In Labrador, comparison between the more complex dynamic fire extension and a classic fire simulator based on a simple fire size distribution showed little difference in terms of mean fire rotation and potential severity, but cumulative burn patterns created by the dynamic fire extension were more heterogeneous due to feedback between fuel types and fire behavior. Simulations in the Sierra Nevada indicated that burn patterns were responsive to topographic features, fuel types, and an extreme weather scenario, although the magnitude of responses depended on elevation. In both study areas, simulated fire size and resulting fire rotation intervals were moderately sensitive to parameters controlling the curvilinear response between fire spread and weather, as well as to the assumptions underlying the correlation between weather conditions and fire duration. Potential fire severity was more variable within the Sierra Nevada landscape and also was more sensitive to the correlation between weather conditions and fire duration. The fire modeling approach described here should be applicable to questions related to climate change and disturbance interactions, particularly within locations characterized by steep topography, where temporally or spatially dynamic vegetation significantly influences spread rates, where fire severity is variable, and where multiple disturbance types of varying severities are common.     

A model for wildfire prevention planning in game resources

The high incidence of hunting activity and forest fires in different ecosystems (particularly in Mediterranean ecosystems) requires a model, which allows for the comprehensive management of hunting resources and estimating the potential damage caused by this type of disturbance. This paper proposes a model for evaluating the socio-economic effects of forest fires on hunting. Its cornerstone lies in evaluating hunting resource availability for each ecosystem within its territorial mosaic. Each game management unit (GMU) is identified by vegetation structure and habitat type. It presents a novel approach in which the economic value of each game management unit is linked to potential forest fire damages. The effect a forest fire has on an entire ecosystem depends on the intensity of its flames. A sample study was undertaken in the province of Huelva in Andalusia (southern Spain). The socio-economic hunting vulnerability of the province of Huelva was 45,188,000€. The results obtained confirmed the need for an economic appraisal of non-forest products in the forest and other woodland areas. Geographic Information System increases the flexibility and simplicity of our methodology which permits immediate extrapolation to other agroforestry territories.     

Trait similarity patterns within grass and grasshopper communities: multitrophic community assembly at work

Trait-based community assembly theory suggests that trait variation among co-occurring species is shaped by two main processes: abiotic filtering, important in stressful environments and promoting similarity, and competition, more important in productive environments and promoting dissimilarity. Previous studies have indeed found trait similarity to decline along productivity gradients. However, these studies have always been done on single trophic levels. Here, we investigated how interactions between trophic levels affect trait similarity patterns along environmental gradients. We propose three hypotheses for the main drivers of trait similarity patterns of plants and herbivores along environmental gradients: (1) environmental control of both, (2) bottom-up control of herbivore trait variation, and (3) top-down control of grass trait variation. To test this, we collected data on the community composition and trait variation of grasses (41 species) and grasshoppers (53 species) in 50 plots in a South African savanna. Structural equation models were used to investigate how the range and spacing of within-community functional trait values of both grasses and their insect herbivores (grasshoppers; Acrididae) respond to (1) rainfall and fire frequency gradients and (2) the trait similarity patterns of the other trophic level. The analyses revealed that traits of co-occurring grasses became more similar toward lower rainfall and higher fire frequency (environmental control), while showing little evidence for top-down control. Grasshopper trait range patterns, on the other hand, were mostly directly driven by vegetation structure and grass trait range patterns (bottom-up control), while environmental factors had mostly indirect effects via plant traits. Our study shows the potential to expand trait-based community assembly theory to include trophic interactions.     

Ten years of land cover change on the California coast detected using Landsat satellite image analysis: Part 2—San Mateo and Santa Cruz counties

Landsat satellite imagery was analyzed to generate a detailed record of 10 years of vegetation disturbance and regrowth for Pacific coastal areas of San Mateo and Santa Cruz Counties. The Landsat Ecosystem Disturbance Adaptive Processing System (LEDAPS) methodology, a transformation of Tasseled-Cap data space, was applied to detected changes in perennial coastal shrubland, woodland, and forest cover from 1999 to 2009. Results showed several principal points of interest, within which extensive contiguous areas of similar LEDAPS vegetation change (either disturbed or restored) were detected. Regrowth of evergreen shrub and tree cover was prevalent along the several long stretches of the coast highway (CA Route 1) between the cities of Half Moon Bay and Santa Cruz. A number of state parks areas showed measurable vegetation restoration as well. The most prominent loss of perennial coastal vegetation over decade was in the Pescadero Marsh area, where the continued presence of levees has historically reduced flood conveyance capacity into and through the marshlands. Based on these examples, the LEDAPS methodology was determined to be capable of fulfilling much of the need for continual, low-cost monitoring of emerging changes to coastal ecosystems.     

Changes in vegetation persistence across global savanna landscapes, 1982–2010

We present a global analysis of the changing face of vegetation persistence in savanna ecosystems by boreal seasons. We utilized nearly 30 years of monthly normalized difference vegetation index data in an innovative time-series approach and developed associated statistical significance tests, making the application of continuous vegetation metrics both more rigorous and more useful to research. We found that 8,000,000–11,000,000 km² of savanna have experienced significant vegetation decline during each season, while 20,000,000–23,000,000 km² have experienced an increase in vegetation persistence during each season, relative to the baseline period (1982–1985). In addition, with the exception of the March–April–May season, which is mixed, the pattern of significant vegetation persistence in the Northern Hemisphere is almost exclusively positive, while it is negative in the Southern Hemisphere. This finding highlights the increasing vulnerability of the Southern Hemisphere savanna landscapes; either resulting from changing precipitation regimes (e.g., southern Africa) or agricultural pressures and conversions (e.g., South America).     

Ten years of land cover change on the California coast detected using landsat satellite image analysis: part 1—Marin and San Francisco counties

Landsat satellite imagery was analyzed to generate a detailed record of 10 years of vegetation disturbance and regrowth for Pacific coastal areas of Marin and San Francisco Counties. The Landsat Ecosystem Disturbance Adaptive Processing System (LEDAPS) methodology, a transformation of Tasseled-Cap data space, was applied to detected changes in perennial coastal shrubland, woodland, and forest cover from 1999 to 2009. Results showed several principal points of interest, within which extensive contiguous areas of similar LEDAPS vegetation change (either disturbed or restored) were detected. Regrowth areas were delineated as burned forest areas in the Point Reyes National Seashore (PRNS) from the 1995 Vision Fire. LEDAPS-detected disturbance patterns on Inverness Ridge, PRNS in areas observed with dieback of tanoak and bay laurel trees was consistent with defoliation by sudden oak death (Phytophthora ramorum). LEDAPS regrowth pixels were detected over much of the predominantly grassland/herbaceous cover of the Olema Valley ranchland near PRNS. Extensive restoration of perennial vegetation cover on Crissy Field, Baker Beach and Lobos Creek dunes in San Francisco was identified. Based on these examples, the LEDAPS methodology will be capable of fulfilling much of the need for continual, low-cost monitoring of emerging changes to coastal ecosystems.     

Response of Savanna Fire Regimes to Changing Fire-Management Policies in a Large African National Park

Abstract:  Approaches to fire management in the savanna ecosystems of the 2-million ha Kruger National Park, South Africa, have changed several times over the past six decades. These approaches have included regular and flexible prescribed burning on fixed areas and a policy that sought to establish a lightning-dominated fire regime. We sought to establish whether changes in management induced the desired variability in fire regimes over a large area. We used a spatial database of information on all fires in the park between 1957 and 2002 to determine elements of the fire regime associated with each management policy. The area that burned in any given year was independent of the management approach and was strongly related to rainfall (and therefore grass fuels) in the preceding 2 years. On the other hand, management did affect the spatial heterogeneity of fires and their seasonal distribution. Heterogeneity was higher at all scales during the era of prescribed burning, compared with the lightning-fire interval. The lightning-fire interval also resulted in a greater proportion (72% vs. 38%) of the area burning in the dry season. Mean fire-return intervals varied between 5.6 and 7.3 years, and variability in fire-return intervals was strongly influenced by the sequencing of annual rainfall rather than by management. The attempt at creating a lightning-dominated fire regime failed because most fires were ignited by humans, and the policy has been replaced by a more pragmatic approach that combines flexible prescribed burning with lightning-ignited fires.