EFFECTS OF FIRE AND HERBIVORY ON THE STABILITY OF SAVANNA ECOSYSTEMS (original) (raw)
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Vegetation, Fire, and Feedbacks: A Disturbance-Mediated Model of Savannas
abstract: Savanna models that are based on recurrent disturbances such as fire result in non-equilibrium savannas, but these models rarely incorporate vegetation feedbacks on fire frequency or include more than two states (grasses and trees). We develop a disturbance model that includes vegetation-fire feedbacks, using a system of differential equations to represent three main components of savannas: grasses, fire-tolerant savanna trees, and fire-intolerant forest trees. We investigate the stability of savannas in the presence of positive feedbacks of fire frequency with (1) grasses, (2) savanna trees, and (3) grasses and savanna trees together while also allowing for negative feedbacks of forest trees on fire frequency. We find that positive feedbacks between fire frequency and savanna trees, alone or together with grasses, can stabilize savannas, blocking the conversion of savannas to forests. Negative feedbacks of forest trees on fire frequency shift the range of parameter space that supports savannas, but they do not generally alter our results. We propose that pyrogenic trees that modify characteristics of fire regimes are ecosystem engineers that facilitate the persistence of savannas, generating both threshold fire frequencies with rapid changes in community composition when these thresholds are crossed and hystereses with bistable community states.
Agriculture, Ecosystems & Environment (Elsevier), 2014
Tropical dry forests and savannas are important repositories of plant diversity and ecosystem services in the tropics. These ecosystems are also used extensively for grazing by livestock, and represent a critical element of the rural economy of many tropical countries. Fire is considered as a part of co-evolution in these ecosystems across the globe. However, in recent decades, there has been a shift in historical fire regime. Fire has become more frequent in these landscapes, and could be further enhanced under climate change. This poses threats to existing biodiversity, ecosystem processes, and rural economy. We asked how variability in fire frequency has influenced diversity and heterogeneity in grass species composition, and richness and abundance of grass species preferred by large herbivores (referred to as grazing accept- ability) in a South Indian tropical savanna forest. We assumed that an increase in fire frequency acts as the active constraint and limits an ecosystem from attaining the maximum heterogeneity, and the maximum grazing acceptability (maximum richness and abundance of grass species preferred by herbivores) in its native settings. We used MODIS active fire and burned area products to estimate fire frequency across the landscape. A nested sampling approach was used to collect information on vegetation and soil at different fire frequencies. Quantile regression analyses indicated that diversity and heterogeneity in grass species composition as well as grazing acceptability decreased with increasing fire frequencies. We found that livestock grazing intervened with the observed vegetation patterns; grass species diversity and hetero- geneity, and grazing acceptability increased with grazing intensity at lower quantiles. Other measured covariates, rainfall, and soil-fertility, alone were not able to explain the observed vegetation patterns in the landscape. The results show a need to control annual fires but allow and manage intermittent fires in this landscape. A complete suppression of fire is not desirable as fire releases nutrients from burning of deeper-rooted vegetation and thus acts as a periodic nutrient pump. It also played an important role in maintaining the grass cover by reducing shrub cover. Hence, it is important to consider the complex interactions between fires–grazers–soil–vegetation to develop effective management practices. We con- clude that fire frequency should be managed at low to intermediate levels (one fire in every 5–9 years, resembling the native settings), and grazing regulated, in order to sustain wild and domestic herbivores, biodiversity, and other key ecosystem processes and services over the long-term.
Tree–grass co-existence in savanna: Interactions of rain and fire
2010
The mechanisms permitting the co-existence of tree and grass in savannas have been a source of contention for many years. The two main classes of explanations involve either competition for resources, or differential sensitivity to disturbances. Published models focus principally on one or the other of these mechanisms. Here we introduce a simple ecohydrologic model of savanna vegetation involving both competition for water, and differential sensitivity of trees and grasses to fire disturbances. We show how the co-existence of trees and grasses in savannas can be simultaneously controlled by rainfall and fire, and how the relative importance of the two factors distinguishes between dry and moist savannas. The stability map allows to predict the changes in vegetation structure along gradients of rainfall and fire disturbances realistically, and to clarify the distinction between climate- and disturbance-dependent ecosystems.
Australian Journal of Botany, 2010
Although the demography of woody plants in savannas has long been shown to be due to many factors, there still is no consensus as to the relative importance of the top-down processes of fire and herbivory, nor on how fire and herbivory affect plant demography. We review the recent literature and suggest that further progress depends on the following: (i) a demographic framework with clear terminology and which focuses on recruitment, transitions and mortality, (ii) an understanding of mechanisms of how fire actually damages plants and how plants survive and out-grow this damage, mainly through height, bark thickness or diameter growth, (iii) an understanding of how losses in biomass due to herbivory may affect plant demography and, (iv) a consideration of interactions between fire and herbivory. Our synthesis suggests (i) strong recruitment limitation as well as some evidence of transition limitation by both fire and herbivory, (ii) that in some cases herbivory alone, notably by elephants and impala, can be more significant than fire alone, on woody plant population size, (iii) that fire and herbivory together are a lethal combination for woody plants and, (iv) that differences in strategies and responses of savanna plants to fire and herbivory are poorly explored.
Trees, grass, and fire in humid savannas-The importance of life history traits and spatial processes
We develop a model to investigate how trees can invade the grass stratum in humid savannas despite repeated fires. In the literature, it is clear that fire reduces tree canopy in savannas. However, fire alone may not be sufficient to prevent tree invasion because there are ecological mechanisms that hamper fire spread by undermining the continuity and density of the grass stratum, which is the means of fire propagation in savannas. Our model is spatially explicit and individual-based, and includes two important factors characterizing the interactions between fire, trees, and grass in savannas, viz. space and the strategies that trees use to cope with fire. The strategies that trees employ against fire emerge from life history traits. According to these strategies, we classify savanna trees into three categories: resprouters, which are able to resprout after their aboveground biomass is burned; resisters, which are able to resist fire due to a thick bark even in the juvenile stages; avoiders, which are very fire-vulnerable in the juvenile stages, but are able to grow fast in the absence of fire. Our results show that trees can invade the grass stratum and finally suppress fire spread because one of the following occurs: (1) trees may resprout and form a population that persists despite repeated effective fires; (2) trees may be fire-resistant; (3) if trees are fire-vulnerable they may cluster and grow in density until fire is prevented. Our results show that fire can be effective in preventing the initiation of the invasion process in the grass stratum. However, once the invasion process has begun, fire alone is not able to reverse this process because of the strategies employed by trees. Furthermore, when a high tree density is reached, grass density is insufficient to allow effective fire spread. From a management point of view, our results imply that fire must be coupled with other factors (browsing, mechanical thinning) to reduce tree density in encroached areas.
Journal of Theoretical Biology, 2011
The tree–grass co-existence in savannas involves multiple and sometimes connected biogeophysical conditions. The savanna domain, its boundaries, and transitions (gradual or abrupt) to other vegetation types (i.e., grassland or forest) are fundamental for the management of ecosystems and for preserving the biodiversity in present conditions and in future changing scenarios. Here we investigate the savanna domain within grazers-fire and browsers-fire parameter planes through a simple ecohydrological model of tree–grass–soil water dynamics. Stability maps allow to identify savanna domains and to show the behavior of vegetation under increasing pressure of grazing and browsing. Stability maps shed light on the causes behind possible vegetation abrupt transitions (e.g., forest collapse and bush encroachment). An application to 15 African savannas sites is presented and discussed with the support of a local sensitivity analysis of the model's parameters.► We investigate the existence and stability of savannas in the herbivores-fire space. ► The analysis is based on a spatially implicit ecohydrological model. ► The results show the types of savannas as function of the disturbance levels. ► An application is discussed with the help of a local sensitivity analysis. ► The comparison between model and data supports the theoretical findings.
Fire
There has been a long-standing interest in understanding how interactions between fire and herbivory influence woody vegetation dynamics in savanna ecosystems. However, controlled, replicated experiments examining how different fire regimes interact with different herbivore groups are rare. We tested the effects of single and repeated burns, crossed with six replicated herbivore treatments, on the mortality and growth of woody vegetation in the Kenya Long-term Exclosure Experiment plots located in a semi-arid savanna system in central Kenya. Burned plots experienced higher tree mortality overall, but differences between burns and non-burns were only significant in plots excluding all wild herbivores and in plots accessible to megaherbivores. Cattle ameliorated the negative effects of repeat burns on tree mortality, perhaps by suppressing fuel load accumulation. Across all herbivore treatments, trees experienced a significant reduction in height within the first two years after fire ...
Effects of four decades of fire manipulation on woody vegetation structure in Savanna
Ecology, 2007
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 d...
International Journal of Wildland Fire
Both wild and prescribed fire in savanna ecosystems influence habitat use by herbivores by creating or maintaining spatial and temporal heterogeneity in forage quality and vegetation cover. Yet little is known about how spatial scales influence long-term persistence of fire effects. We examined changes over a 6-year period in herbivore preference for experimentally burned patches that varied in spatial extent and grain. Avoidance for the burns by elephants and preference for the burns by impala and Grant’s gazelle decreased significantly. For the rest of the species (zebra, eland, oryx, hartebeest, warthog and hare), there were no significant changes in preference for the burns. Changes in preference for the burned areas depended on the spatial extent and grain of the burn, with intermediate-size (9-ha) burns and large (8-ha) patchy burns being more preferred 6–7 years after fire. Grain, but not the spatial extent of the burned area, influenced changes in grass height. Fire resulted...