Woodland dynamics under the influence of elephants and fire in Northern Botswana

Sustained elephant browsing and intense burning could result in the loss of woodlands under conditions where elephant densities are high, such as in northern Botswana. Three woodland types dominated by Acacia erioloba, Baikiaea plurijuga and Colophospermum mopane were monitored in plots and contemporary recruitment rates of woody plants were compared with the associated local elephant densities and fire occurrences. Woodland types differed with respect to structure, extent of elephant damage and the occurrence of fire. Canonical correlations indicated that high extent of fire damage and high elephant densities did not covary within the woodland types investigated. Low tree densities in some woodland types were associated with high elephant densities and new elephant damage to plants increased with high elephant densities during the dry season. Plots with an apparent high fire frequency had lower tree densities and higher cover abundance of shrubs and seedlings.The annual rates of tree recruitment/loss in each woodland type were estimated through a model based on observed seedling recruitment, mortality and reversal to lower height classes due to combinations of fire occurrence and elephant browsing. The model suggested that elephants induce tree loss in woodlands dominated by plant species which are principal food sources. Fire however, seems to have a widespread effect across woodlands which could result in extensive tree loss.     

A Meta-Analysis of the Impact of African Elephants on Savanna Vegetation

Abstract: Large herbivores such as elephants (Loxodonta africana) apparently have a negative impact on woody vegetation at moderate to high population densities. The confounding effects that fire, drought, and management history have may complicate assignment of such impacts to herbivory. We reviewed 238 studies published over 45 years and conducted a meta-analysis based on 21 studies that provided sufficient information on response of woody vegetation to elephants. We considered size and duration of studies, elephant densities, rainfall, fences, and study outcomes in our analysis. We detected a disproportionate citation of 20 published studies in our database, 15 of which concluded that woody vegetation responded negatively to elephants. Our analysis showed that high elephant densities had a negative effect on woody vegetation but that rainfall and presence of fences influenced these effects. In arid savannas, woody vegetation always responded negatively to elephants. In transitional savannas, an increase in elephant densities did not influence woody vegetation response. In mesic savannas, negative responses of woody vegetation increased when elephants occurred at higher densities, whereas elephants confined by fences also had more negative effects on woody plants than elephants that were not confined. Our analysis suggested that rainfall and fences influenced elephant density related impact and that research results were often site-specific. Local environmental conditions and site-specific objectives should be considered when developing management actions to curb elephant impacts on woody vegetation.     

Changes in structure of savanna woodlands in northern Botswana following the impacts of elephants and fire

Fluctuations in densities of woody plant species were monitored in plots within three northern Botswana woodland types subjected to elephant damage and burning. Woodlands dominated by Baikiaea plurijuga and Colophospermum mopane sustained significant changes occurring on an annual basis, whereas Acacia erioloba plots maintained a typical structure. The structure of A. erioloba woodlands appeared to be influenced by factors other than elephants and the occurrence of fire. Woodlands dominated by C. mopane plants were subjected to obtrusive elephant damage, although the densities of tall trees remained largely unchanged. The effects of fire were most prominent in B. plurijuga woodlands. Tree densities declined consistently and plants of lower height classes, such as shrubs and seedlings increased in densities in areas subjected to a high occurrence of fire.     

Humans and elephants as treefall drivers in African savannas

Humans have played a major role in altering savanna structure and function, and growing land‐use pressure will only increase their influence on woody cover. Yet humans are often overlooked as ecological components. Both humans and the African elephant Loxodonta africana alter woody vegetation in savannas through removal of large trees and activities that may increase shrub cover. Interactive effects of both humans and elephants with fire may also alter vegetation structure and composition. Here we capitalize on a macroscale experimental opportunity – brought about by the juxtaposition of an elephant‐mediated landscape, human‐utilized communal harvesting lands and a nature reserve fenced off from both humans and elephants – to investigate the influence of humans and elephants on height‐specific treefall dynamics. We surveyed 6812 ha using repeat, airborne high resolution Light Detection and Ranging (LiDAR) to track the fate of 453 685 tree canopies over two years. Human‐mediated biennial treefall rates were 2–3.5 fold higher than the background treefall rate of 1.5% treefall ha^–1, while elephant‐mediated treefall rates were 5 times higher at 7.6% treefall ha^–1 than the control site. Model predictors of treefall revealed that human or elephant presence was the most important variable, followed by the interaction between geology and fire frequency. Treefall patterns were spatially heterogeneous with elephant‐driven treefall associated with geology and surface water, while human patterns were related to perceived ease of access to wood harvesting areas and settlement expansion. Our results show humans and elephants utilize all height classes of woody vegetation, and that large tree shortages in a heavily utilized communal land has transferred treefall occurrence to shorter vegetation. Elephant‐ and human‐dominated landscapes are tied to interactive effects that may hinder tree seedling survival which, combined with tree loss in the landscape, may compromise woodland sustainability.     

The influence of tree canopies and elephants on sub-canopy vegetation in a savannah

The apparent influence of elephants on the structure of savannahs in Africa may be enhanced by management activities, fire and other herbivores. We separated the effect elephants have on grasses, woody seedlings (<0.5 m) and saplings (0.5–2 m) from the effect of tree canopies (canopy effect), and herbivory (park effect). We defined the canopy effect as the differences between plant abundances and diversity indices under tree canopies and 20 m away from these. Our testing of the park effect relied on the differences in the sub-canopy plant indices inside and outside a protected area that supported a range of herbivores. We based our assessment of the elephant effect on sub-canopy vegetation indices associated with elephant induced reductions in tree canopies. The park and canopy effects were more pronounced than the elephant effect. The park effect suppressed the development of woody seedlings into saplings. Conditions associated with tree canopies benefited woody plants, but not the grasses, as their indices were lower under trees. Elephants reducing canopies facilitated grass species tolerant of direct solar radiation. We concluded that management should consider other agents operating in the system when deciding on reducing the impact that elephants may have on vegetation.     

Changes in the structure and composition of miombo woodlands mediated by elephants (Loxodonta africana) and fire over a 26-year period in north-western Zimbabwe

Changes in structure and composition of miombo woodlands mediated by elephants and fire were studied in 26-year-old permanent transects established in 1972 in north-western Zimbabwe. Elephants caused 48% decline in proportions of large trees (>11 cm diameter), significant reductions (30.9–90.9%) in tree heights, reductions in stem areas (43.5%) and densities (2.5%) of all trees. There were increases in proportions of small trees (64.8%), shrub canopy volumes (271%) and shrub densities (172%). These increases are attributed to natural recruitment because of longer fire-free periods and reduction of tree suppression effects on lower strata as a result of elephant-induced tree declines. Frequencies of occurrence of most species dropped by 28–89.6%. Brachystegia boehmii was replaced by Pseudolachnostylis maprouneifolia as the most dominant tree, largely because of high elephant preference for Brachystegia boehmii . A new suite of species, dominated by Combretaceae, increased in dominance resulting in local floristic changes. Reductions in old elephant (33.4%), old unknown (89.9%) and new elephant (13.7%) damage suggest that elephant occupancy of miombo woodlands has declined, possibly because of limited availability of preferred browse species. This study clearly shows that elephants and fire have contributed significantly to the changes in miombo woodlands in the area.     

Evaluation of the effectiveness of an early peripheral burning strategy in controlling wild fires in north‐western Zimbabwe

A review of the occurrence of wild fires in Sengwa Wildlife Research Area (SWRA), Zimbabwe, is presented for the period 1965–1993. The effectiveness and desirability of early burning of peripheral areas introduced in 1979 are evaluated. More than 75% of wild fires occurred between July and October, 48.6% of which originated from communal lands. Early burning of peripheral areas led to significant reductions in extent of areas burnt, from annual mean areas of 115 km² (1965–1978) to 11 km² (1979–1993) because of effective control of fires, which originated from communal lands. Some areas did not burn at all after 1979, and the overall probability of burning dropped from 0.484 to 0.187. Whilst it may be desirable to keep fires out of SWRA in the short term, fuel build‐ups increase the fire hazard resulting in negative consequences on biodiversity in the long term. Consideration should be given to combine peripheral burning with low‐intensity prescribed burning of selected blocks to keep a semblance of natural fire regimes to ensure the maintenance of biodiversity while simultaneously reducing the fire‐hazard. An integrated fire management plan should be put in place for SWRA.     

Woody cover in African savannas: the role of resources, fire and herbivory

Aim To determine the functional relationships between, and the relative importance of, different driver variables (mean annual precipitation, soil properties, fire and herbivory) in regulating woody plant cover across broad environmental gradients in African savannas. Location Savanna grasslands of East, West and Southern Africa. Methods The dependence of woody cover on mean annual precipitation (MAP), soil properties (texture, nitrogen mineralization potential and total phosphorus), fire regimes, and herbivory (grazer, browser + mixed feeder, and elephant biomass) was determined for 161 savanna sites across Africa using stochastic gradient boosting, a refinement of the regression tree analysis technique. Results All variables were significant predictors of woody cover, collectively explaining 71% of the variance in our data set. However, their relative importance as regulators of woody cover varied. MAP was the most important predictor, followed by fire return periods, soil characteristics and herbivory regimes. Woody cover showed a strong positive dependence on MAP between 200 and 700 mm, but no dependence on MAP above this threshold when the effects of other predictors were accounted for. Fires served to reduce woody cover below rainfall‐determined levels. Woody cover showed a complex, non‐linear relationship with total soil phosphorus, and was negatively correlated with clay content. There was a strong negative dependence of woody cover on soil nitrogen (N) availability, suggesting that increased N‐deposition may cause shifts in savannas towards more grassy states. Elephants, mixed feeders and browsers had negative effects on woody cover. Grazers, on the other hand, depressed woody cover at low biomass, but favoured woody vegetation when their biomass exceeded a certain threshold. Main conclusions Our results indicate complex and contrasting relationships between woody cover, rainfall, soil properties and disturbance regimes in savannas, and suggest that future environmental changes such as altered precipitation regimes, N‐enrichment and elevated levels of CO₂ are likely to have opposing, and potentially interacting, influences on the tree–grass balance in savannas.     

Predicting the Effects of Woody Encroachment on Mammal Communities,Grazing Biomass and Fire Frequency in African Savannas

With grasslands and savannas covering 20% of the world’s land surface, accounting for 30–35% of worldwide Net Primary Productivity and supporting hundreds of millions of people, predicting changes in tree/grass systems is priority. Inappropriate land management and rising atmospheric CO₂ levels result in increased woody cover in savannas. Although woody encroachment occurs world-wide, Africa’s tourism and livestock grazing industries may be particularly vulnerable. Forecasts of responses of African wildlife and available grazing biomass to increases in woody cover are thus urgently needed. These predictions are hard to make due to non-linear responses and poorly understood feedback mechanisms between woody cover and other ecological responders, problems further amplified by the lack of long-term and large-scale datasets. We propose that a space-for-time analysis along an existing woody cover gradient overcomes some of these forecasting problems. Here we show, using an existing woody cover gradient (0–65%) across the Kruger National Park, South Africa, that increased woody cover is associated with (i) changed herbivore assemblage composition, (ii) reduced grass biomass, and (iii) reduced fire frequency. Furthermore, although increased woody cover is associated with reduced livestock production, we found indigenous herbivore biomass (excluding elephants) remains unchanged between 20–65% woody cover. This is due to a significant reorganization in the herbivore assemblage composition, mostly as a result of meso-grazers being substituted by browsers at increasing woody cover. Our results suggest that woody encroachment will have cascading consequences for Africa’s grazing systems, fire regimes and iconic wildlife. These effects will pose challenges and require adaptation of livelihoods and industries dependent on conditions currently prevailing.     

Seronera bull problem: The trees

(1) The impact of groups of bull elephants on the wooded–grassland of the Seronera area of the Serengeti National Park, Tanzania, is examined. (2) Elephant use of trees occurs contagiously. At use loci in the woodlands, removal of canopy–forming acacia has been as high as 6% per annum. (3) However, from a survey of the whole Seronera woodland community, the loss of large trees has been 2–5% per annum. (4) It is argued that the regeneration potential is adequate to compensate loss due to elephant activity, and that large acacia will not disappear because of elephants. (5) Management action is discussed and recommendations made.     

Trends in woody vegetation cover in the Kruger National Park, South Africa, between 1940 and 1998

Changes in the cover and density of shrubs and trees were assessed from aerial photographs (1940, 1974 and 1998) as well as from fixed-point photographs taken in 1984 and 1996 in the Kruger National Park, South Africa. Woody cover (trees and shrubs combined) increased by 12% on granite substrates but decreased by 64% on basalt substrates over the past 58 years. Both these figures are expressed in terms of the initial values, respectively. The density of the large tree component of woody vegetation decreased on both substrates. Woody vegetation cover declined as fire return periods became shorter, but the relationship was weak. The increases in woody plant density and cover on granite are thought to be the result of decreased competition from grasses, which in turn is a result of overgrazing by wild herbivores whose numbers have been kept high through the provision of surface water. These effects were not seen on the relatively nutrient-rich basalts, where grasses can recover rapidly even after heavy grazing. The decline in overall woody cover on basalts is interpreted as a result of regular, short-interval prescribed burning over the past 40 years, while the universal decline in large trees seems to result from an interaction between regular, frequent fires and utilization by elephants. The implications for management are discussed.     

Elephant herbivory in Majete Wildlife Reserve,Malawi

Elephants have a major influence on vegetation structure, composition and ecosystem processes, and are primary agents of habitat change in Africa. At moderate‐to‐high population densities, elephants can damage vegetation, especially when enclosed in protected areas. This study examines the effects of elephant browsing on woody trees in Majete Wildlife Reserve (WR), south‐western Malawi. Regression analysis is used to assess the associations of six factors known to drive elephant browsing in other areas and determine which ones have the most influence on browsing at Majete WR. Twenty‐four per cent of tagged trees had been subject to elephant browsing. The model with vegetation type, stem diameter and distance from permanent water correctly predicted browsing for 80% of the observations. Elephants mostly favoured riparian woodlands, followed by Acacia‐dominated woodland and Brachystegia‐dominated woodland. Browsing occurrence was negatively related to distance from permanent water and diameter at breast height(DBH). A larger number of trees, sampled at random and covering a larger portion of the reserve would provide more reliable estimates of browsing and related factors. Knowledge of time‐ and site‐specific factors affecting elephant browsing can be used to forecast future habitat transformations and manipulate the range of the elephants within the reserve.     

Precipitation and elephants,not fire,shape tree community composition in Serengeti National Park,Tanzania

Savannas are spatially diverse, variable and are susceptible to high rates of disturbance from fire and herbivory. There is significant interest in woody cover dynamics in relation to disturbance regimes. Less effort has been devoted to understand processes that drive tree community composition. In this study, tree species composition data collected at the landscape scale in the Serengeti were used to identify key environmental factors driving variation in species composition. A system of 38 plots clustered within 10 sites spanning the mean annual precipitation (MAP) gradient was used to assess the relative role of bottom‐up (precipitation, soil nutrients and soil texture) vs. top‐down factors (fire and elephant herbivory) on tree community composition. We developed candidate models relating tree species composition (based on multivariate community analysis) to different combinations of plot‐level environmental covariates. Results suggest that tree community composition is largely driven by MAP and is associated with elephant population density. Strikingly, we found no evidence that fire influences species compositional turnover. In a second analysis, we used structural equation model (SEM) to explore the possible direction of association between elephant density and tree species composition. We compared a model that included elephant effects on composition to one that included community composition effects on elephant density. Results suggest that variation in elephant population density across space is more likely to drive tree community composition and not vice versa. We propose that precipitation and herbivory, rather than fire, determine tree species composition in Serengeti Acacia tree community.     

Long-term patterns of shrub expansion in a C4-dominated grassland: fire frequency and the dynamics of shrub cover and abundance

Worldwide, grassland ecosystems have experienced a major shift in growth-form dominance as woody plant species have expanded and replaced native grasses. In the C(4)-dominated grasslands of central North America, a reduction in fire frequency is the most cited cause of this shift in growth forms as fire both enhances grass productivity and constrains the establishment and expansion of native woody vegetation. Using an 18-yr plant species composition data set, we quantified patterns of change in shrub cover, frequency, and species richness associated with three distinct fire regimes. During the study period (1983-2000), shrub cover increased most dramatically in sites in which the frequency of fire was once every 4 yr (intermediate frequency; 28.6%) followed by sites in which fire occurred only once during the 18-yr period (low frequency; 23.7%). Annual fire effectively prevented the recruitment of new woody species, but even with this high fire frequency, shrub cover increased slightly (3.7%). Comparatively, shrub species richness increased by three and six, respectively, in the intermediate- and low-frequency fire sites. These data indicate that within this grassland, periods without fire are necessary for recruitment of both new individuals and additional shrub species; however, once established, shrub cover will increase regardless of fire frequency and even annual fire will not reduce shrub abundance.     

Growth response of woody species to elephant foraging in Mwea National Reserve,Kenya

The African elephant (Loxodonta africana) is known to greatly affect the structure and dynamics of vegetation. In Mwea National Reserve, elephants foraged mainly on Acacia ataxacantha and Grewia bicolor out of the five most preferred woody species. However, out of the five preferred woody species, only Grewia virosa and G. bicolor showed a positive association between their fresh use and past elephant use. All the five selected woody species showed high coppicing response after foraging, with the highest coppice growth rates recorded for Acacia brevispica and lowest for Grewia tembensis. The mean heights of woody species utilized by elephants were highest for A. brevispica and lowest for G. bicolor. The mean heights of coppices emerging after utilization by elephants were not significantly different for A. ataxacantha but were significantly shorter in the rest of the foraged species. Elephants avoided the coppices of many other woody species notably C. africana, A. tortilis, A. mellifera, Combretum aculeatum among others in the reserve. The objective of this study was to understand the capacity of woody species to recover after utilization by elephants and feeding response of elephants to new woody species re‐growth; a cycle that would define the dynamics of food resources and elephant population within the reserve.     

Elephant survival,rainfall and the confounding effects of water provision and fences

Elephant are increasing across some areas of Africa leading to concerns that they may reduce woodlands through their feeding. Droughts may help limit elephant numbers, but they are generally both episodic and local. To explore more general impacts of rainfall, we examine how its annual variation influences elephant survival across ten sites. These sites span an almost coast-to-coast transect of southern Africa that holds the majority of the ~500,000 remaining savanna elephants. Elephants born in high rainfall years survive better than elephants born in low rainfall years. The relationship is generally weak, except at the two fenced sites, where rainfall greatly influenced juvenile survival. In these two sites, there are also extensive networks of artificial water. Rainfall likely affects elephant survival through its influence on food. The provision of artificial water opens new areas for elephants in the dry season, while fencing restricts their movements in the wet season. We conclude that the combination of these factors makes elephant survival more susceptible to reductions in rainfall. As a result, elephants living in enclosed reserves may be the first populations to feel the impacts of global warming which will decrease average rainfall and increase the frequency of droughts. A way to prevent these elephants from damaging the vegetation within these enclosed parks is for managers to reduce artificial water sources or, whenever practical, to remove fences.     

Elephant spatial use in wet and dry savannas of southern Africa

The influence of elephants on woody vegetation cover varies from place to place. In part this may be due to the way elephants utilize space across landscapes and within their home ranges in response to the availability and distribution of food. We used location data from 18 cows at six study sites across an east to west rainfall gradient in southern Africa to test whether wet- and dry-season home-range sizes, evenness of space use within seasonal home ranges and range overlap between seasons and between years, differed between wet and dry savannas. We then tested whether the quantity, distribution and seasonal stability in vegetation productivity, a coarse measure of food for elephants, explained differences. Elephants in wet savannas had smaller wet- and dry-season home ranges and also returned to a higher proportion of previously visited grid cells between seasons and between years than elephants living in dry savannas. Wet-season home-range sizes were explained by seasonal vegetation productivity while dry-season home-range sizes were explained by heterogeneity in the distribution of vegetation productivity. The influence of the latter on dry-season home ranges differed among structural vegetation classes. Range overlap between seasons and between years was related to inter-seasonal and inter-annual stability in vegetation productivity, respectively. Evenness of elephant spatial use within home ranges did not differ between savanna types, but it was explained by seasonal vegetation productivity and heterogeneity in the distribution of vegetation productivity during the wet season. Differences in elephant spatial use patterns between wet and dry savannas according to vegetation structure and season may need to be included in the development of site-specific objectives and management approaches for African elephants.     

An elephantine challenge: human–elephant conflict distribution in the largest Asian elephant population,southern India

Wildlife conservation is a complex issue especially when it involves large carnivores or mega-herbivores that are conflict-prone. Karnataka state in southern India is known to harbor high density of wild elephants. This conservation success story also has opportunity costs for communities living in close proximity to elephants. Despite the fact that human–elephant conflict is a serious conservation and social issue, there is little quantitative understanding of conflict especially over large areas. Here we conduct the first analysis of human–elephant conflict distribution, severity and explanatory factors over the entire state of Karnataka. We use data from the state forest department records on villages that experience conflict, compensation payments made by the government, elephant densities, forest cover and perimeter, and presence of physical barriers to mitigate elephant conflict. In total, 60,939 incidences of crop loss were reported and US$ 2.99 m paid in compensation during April 2008–March 2011. A total of 91 people were killed by elephants and 101 elephants died in retaliatory killings during the study period. A total of 9.4 % of the state’s geographic area covering 25 of the 42 forest administrative divisions were affected. There was no significant difference in conflict incidences or compensation given between protected areas and non-protected areas. There was no correlation between conflict incidences/unit area and elephant density, forest cover, forest perimeter of protected areas and presence of physical barriers. The results depict the importance of efficient management of physical barriers, conserving key habitat linkages, and acts as baseline data for future work.     

Structure of population,grouping patterns and density of forest elephants in north-west Congo

The structure of the population of forest elephants visiting a clearing in north-west Congo was studied over 8 months. Out of 3314 sightings, 629 elephants were identified (including 64%, of adults). The sex ratio was about 1:1. The number of offspring per female was 1.12. Solitary elephants (91.4%, of males) made up 35% of the population. Of females, 93.6% were grouped. Groups (mean size 3.5) included females and offspring (3.4), males and females (5.0), or males only (2.3). The return rate of elephants at the clearing suggested that up to 1900 individuals have visited the clearing. This confirms that elephant densities are especially high in north Congo.     

Ants,fire, and bark traits affect how African savanna trees recover following damage

Bark damage resulting from elephant feeding is common in African savanna trees with subsequent interactions with fire, insects, and other pathogens often resulting in tree mortality. Yet, surprisingly little is known about how savanna trees respond to bark damage. We addressed this by investigating how the inner bark of marula (Sclerocarya birrea), a widespread tree species favoured by elephants, recovers after bark damage. We used a long‐term fire experiment in the Kruger National Park to measure bark recovery with and without fire. At 24 months post‐damage, mean wound closure was 98, 92, and 72%, respectively, in annual and biennial burns and fire‐exclusion treatments. Fire exclusion resulted in higher rates of ant colonization of bark wounds, and such ant colonization resulted in significantly lower bark recovery. We also investigated how ten common savanna tree species respond to bark damage and tested for relationships between bark damage, bark recovery, and bark traits while accounting for phylogeny. We found phylogenetic signal in bark dry matter content, bark N and bark P, but not in bark thickness. Bark recovery and damage was highest in species which had thick moist inner bark and low wood densities (Anacardiaceae), intermediate in species which had moderate inner bark thickness and wood densities (Fabaceae) and lowest in species which had thin inner bark and high wood densities (Combretaceae). Elephants prefer species with thick, moist inner bark, traits that also appear to result in faster recovery rates.