Short‐term Effects of Single Species Browsing Release by Different‐sized Herbivores on Sand Forest Vegetation Community,South Africa

Manipulations of herbivores in protected areas may have profound effects on ecosystems. We examine short‐term effects on tree species assemblages and resource utilization by a mesoherbivore and small‐size herbivores (ungulates <20 kg) in Sand Forest, after browsing release from a megaherbivore (elephant), or both a mega‐ and mesoherbivore (nyala), respectively. Effects were experimentally separated using replicated exclosures where all trees were counted, identified to species and browsing events recorded. Tree species assemblages were impacted by both elephant and nyala, and by each herbivore species individually. Tree turnover rates were higher where both herbivore species were present than in their combined absence. Diet was segregated among elephant, nyala and small‐size herbivores. Both resource specificity and browsing pressure by nyala increased in absence of elephant; small‐size herbivores increased resource specificity in absence of elephant, and increased browsing pressure in absence of both elephant and nyala. This implies interference competition with competitive release. The indirect effect of the manipulation of herbivore populations, through the removal of one or two herbivore species, caused a shift in tree species composition and diet of smaller‐size herbivores. These indirect effects, especially on tree species composition, can become critical as they affect vegetation dynamics, biodiversity and ecosystem processes. Therefore, in order to conserve habitats and biodiversity across all trophic levels, conservation managers should consider the effects of: (1) the full herbivore assemblage present; and (2) any effects of altering the relative and absolute abundance of different herbivore species on other herbivore species and vegetation.     

Hierarchical foraging by giraffe in a heterogeneous savannah,Tanzania

Understanding foraging decisions made by wildlife at different spatio‐temporal scales is important for wildlife management and conservation. We tested whether foraging decisions by Masai giraffe (Giraffa camelopardalis tippelskirchi Matschie) differed with scales; habitat, patch and tree in a heterogeneous savannah. We collected data from Arusha National Park, Tanzania, in March–May and August–October 2013. Visual observations were used to collect data on foraging behaviour. Measurements of tree height and stem height and scoring of accumulated browsing were made in 133 patches around trees where the giraffes had been seen browsing, and in a corresponding number of available patches. Giraffes selected Acacia shrub and Dodonea shrub habitats but not for water availability or predator avoidance. For patch use, giraffe selected high quantity of Acacia xanthophloea and Dodonea viscosa. Between plant species, A. xanthophloea was the most preferred and within plant species, tree quality was enhanced by tree height and high score of accumulated browsing. Generally, giraffes selected for A. xanthophloea at all scales.     

The impact of giraffe, rhino and elephant on the habitat of a black rhino sanctuary in Kenya

The habitat in an enclosed black rhino sanctuary, the Sweetwaters Game Reserve in Kenya, is being altered as populations of elephant, giraffe and black rhino increase. Height‐specific browse impact data were recorded for 1075 trees of the dominant species, the whistling thorn, Acacia drepanolobium. Rhinos and elephants browsed 18% of these trees in 1 year, including 5% that were killed or removed. The remaining trees were subjected to high levels of giraffe browse and low rainfall and grew by only 7.5 cm in a year. A mathematical model has been constructed that predicts how the number of trees ha^?1 will change with time under different browsing impacts. The model compares recruitment rate with removal rate and estimates that the number of trees ha^?1 will fall by 2% per year under the current browsing impact of black rhino (0.27 per km²), elephant (1.1 per km²) and giraffe (1.9 per km²). In 7 years, if the rhino and elephant populations continue to increase at the current rates, tree density will be falling by 5% per year and nearly one‐third of the trees will have been removed. These conditions are unsustainable and will result in habitat change and may affect rhino breeding. Several ways of alleviating the problem are discussed.     

Introduction of giraffe changes acacia distribution in a South African savanna

Large mammal herbivores can have significant effects on the structure and composition of plant communities. We studied the impacts of an introduced giraffe population on Acacia species at Ithala Game Reserve in South Africa. Browse intensity and Acacia mortality were assessed in field transects, and in road transects covering the reserve road network. Several Acacia species occurring in high‐density giraffe areas had high levels of mortality. Populations of Acacia davyi were extinct in areas accessible to giraffe. Most A. caffra trees within giraffe browse height were dead and A. karroo, the most common species, was also heavily affected. Some species, including A. tortilis, showed no or very low mortality attributable to giraffe browsing. Healthy populations of sensitive species occurred in areas within, and adjacent to, the reserve in areas with low or no giraffe browsing. Areas too steep for giraffe access formed spatial refuges for these trees. The differential mortality that is occurring as a consequence of giraffe browsing is altering species composition and species distribution in this savanna landscape.     

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.     

Ecological separation among browsing ungulates in Tsavo East National Park,Kenya

Summary Data on food habits and habitat preferences of four browsing herbivores (black rhinoceros, giraffe, gerenuk and lesser kudu) were analyzed to assess niche width for each species and niche overlap between pairs of species. All four species depended heavily on woody plants as food, and overlap in the utilization of different plant types (trees and shrubs, herbs, grasses, etc.) was very great in three of six species paris. When individual plant species were considered, markedly less overlap was apparent. Three of the four ungulate species preferred the most densely wooded vegetation type. Overlap in habitat preferences tended to be least in those pairs of species with the greates dietary overlap, which resulted in some degree of ecological separation. This was further increased by differences in browsing level. Seasonal variations in the browsing level of the giraffe had the effect of reducing overlap with the other species in the dry season, when food was in relatively short supply. Whether or not actual competition existed among the four ungulate species could not be established; in any event, it would probably be less important than possible competition exerted by the elephant, the dominant herbivore by far in the ecosystem. The ecological separation evident among the four browsing species probably permitted them to coexist in the area before the elephant reached its present dominant position and started altering the original vegetation.Formerly Tsavo East Research Station, Voi, Kenya     

Not all ants are equal: obligate acacia ants provide different levels of protection against mega‐herbivores

In obligate ant–plant mutualisms, the asymmetric engagement of a single plant species with multiple ant species provides the opportunity for partners to vary in their behaviour. Variation in behaviour has implications for the interactions with third‐party species such as herbivores. This study assessed the effect of obligate ant‐mutualists (Crematogaster mimosae, Crematogaster nigriceps and Tetraponera penzigi) inhabiting the African ant‐acacia (Acacia drepanolobium) on three mega‐herbivore browsers: the Maasai giraffe (Giraffa camelopardalis tippelskirchi), the reticulated giraffe (Giraffa c. reticulata) and the black rhino (Diceros bicornis). Giraffes are abundant and wide‐ranging herbivores of the acacias, whereas black rhinos are localized and perennial herbivores of the acacias. Multiyear field studies comparing the ants’ aggressive behaviour and browsing by mega‐herbivores suggested differences between the tending abilities of the primary ant species inhabiting A. drepanolobium. Trees occupied by the aggressive ant species C. mimosae had significantly less browsing by giraffes and black rhino than trees occupied by other ant species. The results of this study provide evidence that ant‐mutualists on African acacias can serve as deterrents to mega‐herbivores and that different ant species vary in their tending abilities.     

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.     

Associational refuge in practice: can existing vegetation facilitate woodland restoration?

Herbivores can dramatically diminish revegetation success, but associational refuge theory predicts that neighbouring plants could hinder browsing of planted seedlings. The key to strategic restoration using associational refuge is to define which patch variables are effective against the appropriate herbivores, at multiple scales, and to understand which stages of the foraging process these variables disrupt. Our study aimed to test the capacity of existing vegetation to act as associational refuge for planted seedlings by affecting search, detection and consumption decisions, and more generally influence herbivore foraging patterns. We conducted a field trial with free‐ranging, mammalian herbivores and nursery‐raised, native tree seedlings. We quantified seedling browsing damage over time in relation to a suite of existing patch variables at two spatial scales (100 m² and 4 m²). After two months, 78% of seedlings were browsed, suffering mean foliage loss of 90.5%. Focal seedlings were almost exclusively consumed by swamp wallabies Wallabia bicolor, an abundant generalist browser. Once a swamp wallaby investigated a seedling, the probability of consumption was high (86%). At the large scale, browsing of seedlings was delayed in patches with lower canopy cover and fewer browsed plant species. Seedlings in fern‐dominated patches escaped browsing for longer than those in grass‐dominated patches. At the small scale, browsing was delayed with higher cover of understorey vegetation. Associational refuge was provided by vegetation with characteristics, and at spatial scales, consistent with disrupted search and detection of focal seedlings by herbivores. Thus strategic placement of seedlings in existing vegetation – based on understanding which herbivore species is responsible and how it responds to vegetation – can take advantage of associational refuge during restoration. However, given rapid seedling detection by herbivores, associational refuge may be inadequate in the long‐term under high browsing pressure unless high absolute numbers of seedlings are planted among refuge.     

The savanna ecology of Kidepo Valley National Park

(1) Food habits of elephant and giraffe were recorded in short sample periods over several years using the feeding minutes technique. (2) Twelve species of common plants were collected on these occasions, separated into their main parts and analysed chemically. (3) Damage to trees was also recorded at intervals. (4) Fifty-nine species of plant were eaten by elephant. During wet months their diet was predominantly grass but during dry months woody vegetation became more important. Giraffe were observed eating from thirty-nine species of plant which were almost entirely browse throughout the year. At least eighteen of these species were also eaten by elephant. (5) Three of the browse species most important to giraffe were also important to elephant in dry seasons. In the driest month the species eaten by giraffe formed more than two-thirds of the observed diet of elephants. (6) Chemical analyses of plants show that browse leaves had a higher level of protein, fat and minerals than most other plants. (7) The trunk of an elephant and the tongue of a giraffe are highly sensitive organs which permit food selection. Elephant in particular appeared to alter their food habits seasonally and it is suggested that this may be an attempt to obtain a balanced intake of nutrients. (8) Due to the effect of elephant and other agencies there is an accelerating decline in Acacia gerrardii woodlands. Browsing and fire are independently preventing their regeneration. (9) The extensive overlap of food requirements on preferred but diminishing browse resources implies potential if not already existing competition between elephant and giraffe. (See Part III.)     

Change in Mesoherbivore Browsing Is Mediated by Elephant and Hillslope Position

Elephant are considered major drivers of ecosystems, but their effects within small-scale landscape features and on other herbivores still remain unclear. Elephant impact on vegetation has been widely studied in areas where elephant have been present for many years. We therefore examined the combined effect of short-term elephant presence (< 4 years) and hillslope position on tree species assemblages, resource availability, browsing intensity and soil properties. Short-term elephant presence did not affect woody species assemblages, but did affect height distribution, with greater sapling densities in elephant access areas. Overall tree and stem densities were also not affected by elephant. By contrast, slope position affected woody species assemblages, but not height distributions and densities. Variation in species assemblages was statistically best explained by levels of total cations, Zinc, sand and clay. Although elephant and mesoherbivore browsing intensities were unaffected by slope position, we found lower mesoherbivore browsing intensity on crests with high elephant browsing intensity. Thus, elephant appear to indirectly facilitate the survival of saplings, via the displacement of mesoherbivores, providing a window of opportunity for saplings to grow into taller trees. In the short-term, effects of elephant can be minor and in the opposite direction of expectation. In addition, such behavioural displacement promotes recruitment of saplings into larger height classes. The interaction between slope position and elephant effect found here is in contrast with other studies, and illustrates the importance of examining ecosystem complexity as a function of variation in species presence and topography. The absence of a direct effect of elephant on vegetation, but the presence of an effect on mesoherbivore browsing, is relevant for conservation areas especially where both herbivore groups are actively managed.     

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.     

Spatial dynamics of lions and their prey along an environmental gradient

Establishing the ecological determinants of the spatial dynamics of large African savanna mammals is necessary for understanding the cumulative impacts on the suitability and resilience of their natural habitats, of progressive habitat fragmentation and temporal shifts in climate, fire regimes and elephant browsing. Systematic directional gradients and small‐scale spatial dependence were evident in rainfall, herbivore biomass and lion density in the Masai Mara National Reserve during September 1990 to July 1992. Lion density, the biomass of resident and all herbivore species were autocorrelated within 4.3, 4.1 and 3.3 km, respectively, and peaked in areas of relatively low rainfall but higher microtopographic and vegetation heterogeneity and complexity and more diverse drainage systems. Rainfall influenced herbivore biomass and hence lion density non‐linearly and that influence was apparently modified by forage mineral nutrients, structural habitat complexity, degree of drainage and distance to waterpoints. Lion density was related to the biomass of resident herbivores, even after controlling for rainfall and spatial trends. Persistence of the recent decline in Mara woodlands and increasing human population along the reserve–ranch boundary would likely reduce the reserve's lion population because most lions spent the daytime within the vegetation mosaic fringing drainage lines, away from humans and domestic livestock.     

Female tannin avoidance: a possible explanation for habitat and dietary segregation of giraffes (Giraffa camelopardalis peralta) in Niger

Niger is host to the last free‐roaming herd of G.c. peralta (Giraffa camelopardalis peralta). We examined the foraging preferences of these giraffe in their dry‐season habitats, with the goal of preserving the herd in the regions that they currently inhabit. The current dry‐season habitat comprises two distinct vegetation zones. In both of these zones the giraffe must exist alongside the people of this region. The giraffes exhibit a sexual segregation in their dry‐season habitat selection and forage choices. The females show a strong preference for the intermediate zone (IM) when lactating. The males and pregnant females show a preference for the Dallol Bosso (DB). Nursing cows exhibit an avoidance of tannins. Bulls and non‐nursing cows prefer high protein and high fat forage, while subadults show a strong preference for high protein and carbohydrate contents and moderate tannin levels. Combretum glutinosum is the preferred species for adults of both sexes in the IM. Males and females have strong preferences for both Acacia nilotica and Acacia seyal in the DB. Sub‐adults of both sexes strongly prefer Prosopis africana in the IM. Unlike females, males retain their preference for A. nilotica when in the IM.     

Woody vegetation damage by the African elephant during severe drought at Pongola Game Reserve,South Africa

Elephants (Loxodonta africana) significantly alter ecosystem structure and composition through browsing (e.g. pollarding, debarking and toppling). Such browsing is predicted to intensify during severe drought which may become more common with climate change. Here, we make use of an elephant impact survey from 2012 to 2015 and during the El Nino drought of 2015–2016 at Pongola Game Reserve (107 km²), KwaZulu-Natal, to investigate how severe drought influenced damage severity of different tree heights and species by elephants in this small reserve. Contrary to expectations, damage to common species did not change with severe drought. Crown damage had the highest predicted probability across heights (29%–90%) and species (46%–75%) regardless of drought. However, we found severe drought increased the predicted probabilities of crown damage to smaller trees <4 m, mortality >6 m and severe damage at 4–6 m. Consequently, elephant damage during severe drought may alter vegetation structure by severely damaging or killing large trees (>4 m) and extensively damaging the crowns of trees <4 m. Long-term monitoring of elephant effects on woody vegetation is essential to enable science-based management in response to future drought and elephant damage (e.g. range expansion, beehive deterrents) to protect elephants and conserve woody vegetation.     

Mutualistic acacia ants exhibit reduced aggression and more frequent off‐tree movements near termite mounds

In many ant–plant mutualisms, ants establish colonies in hollow thorns, leaf pouches, or other specialized structures on their host plants, which they then defend from herbivores. Resource heterogeneity could affect the maintenance of these mutualisms if it leads to one or both partners altering their investment in the interaction. Such a phenomenon may be especially pertinent to the Acacia–ant mutualism found in East African savannas, where termite mounds have a profound effect on the spatial structuring of resources used by both plants and ants. Here, we examined whether the proximity to termite mounds of Acacia drepanolobium trees is associated with variation in the behavior of one of their ant associates, Crematogaster nigriceps. We found that ant colonies near termite mounds had decreased aggressive responses to simulated herbivory as well as increased off‐tree movement. We hypothesize that these changes are the result of resident ant colonies near termite mounds shifting investment from defense of their host plant to foraging for nearby resources.     

Grasses and browsers reinforce landscape heterogeneity by excluding trees from ecosystem hotspots

Spatial heterogeneity in woody cover affects biodiversity and ecosystem function, and may be particularly influential in savanna ecosystems. Browsing and interactions with herbaceous plants can create and maintain heterogeneity in woody cover, but the relative importance of these drivers remains unclear, especially when considered across multiple edaphic contexts. In African savannas, abandoned temporary livestock corrals (bomas) develop into long-term, nutrient-rich ecosystem hotspots with unique vegetation. In central Kenya, abandoned corral sites persist for decades as treeless ‘glades’ in a wooded matrix. Though glades are treeless, areas between adjacent glades have higher tree densities than the background savanna or areas near isolated glades. The mechanisms maintaining these distinctive woody cover patterns remain unclear. We asked whether browsing or interactions with herbaceous plants help to maintain landscape heterogeneity by differentially impacting young trees in different locations. We planted the mono-dominant tree species (Acacia drepanolobium) in four locations: inside glades, far from glades, at edges of isolated glades and at edges between adjacent glades. Within each location, we assessed the separate and combined effects of herbivore exclusion (caging) and herbaceous plant removal (clearing) on tree survival and growth. Both caging and clearing improved tree survival and growth inside glades. When herbaceous plants were removed, trees inside glades grew more than trees in other locations, suggesting that glade soils were favorable for tree growth. Different types of glade edges (isolated vs. non-isolated) did not have significantly different impacts on tree performance. This represents one of the first field-based experiments testing the separate and interactive effects of browsing, grass competition and edaphic context on savanna tree performance. Our findings suggest that, by excluding trees from otherwise favorable sites, both herbaceous plants and herbivores help to maintain functionally important landscape heterogeneity in African savannas.     

Facilitation by herbivore-mediated nurse plants in a threatened tree, Taxus baccata: local effects and landscape level consistency

We investigated the regeneration of a threatened tree, the yew Taxus baccata, in relation to the presence of fleshy‐fruited woody plants acting as seed dispersal foci as well as protecting yew recruits against ungulate herbivores. We seek to determine if local facilitative effects are consistent across landscape in the Cantabrian range (NW Spain). Yew seed rain by birds mostly concentrated under yew trees and beneath hollies Ilex aquifolium. Seedling emergence distributed similarly to seed rain, but first‐year seedling survival was higher beneath hollies. In one site where woody vegetation was structured as nucleation centres (multispecific patches of fleshy‐fruited plants acting as foci for seed rain) yew recruits mostly occurred in yew‐dominated centres, suggesting dispersers‐mediated facilitation. However, holly was the main nurse plant for most of these recruits, considering the nurse as the species whose canopy covered directly the yew recruit. Living beneath nurse plants reduced herbivore damage on saplings and enhanced seedling survival. A planting experiment with yew rooted‐cuttings beneath different spiny shrubs corroborated this effect. Additional evidence on yew recruitment limitation by herbivory emerged from one population where ungulates were fence‐excluded. Our results suggest that nurse plants mitigate the negative effect of herbivores on yew regeneration, by providing defence against browsing and trampling. Shelter ability related to nurse structure, cone‐shaped shrubs with branches at their bases acting better as a barrier. Paradoxically, this structure resulted from heavy browsing on nurse plants. The study of yew regeneration and habitat structure in seven sites provided evidence for the consistency of facilitation by holly at the landscape level, since local values of yew recruitment positively related to nurse ground cover. Range‐scale yew management must consider the local functioning of the interaction among avian seed‐dispersers, nurse fleshy‐fruited plants and ungulate herbivores, in combination with regional measures, targeting the habitats where facilitation emerges.     

Food habits of giraffe in Tsavo National Park, Kenya

In the context of a broader ecological investigation, food habits of giraffe were studied in Tsavo National Park (East). The only method employed was direct observation of feeding animals in the field. Each instance in which one animal fed on one individual plant was counted as one food record for this plant species; 4025 records are analysed. A total of sixty-six plant species was found to be eaten, the great majority being trees and shrubs, with a few creepers and vines. There were marked seasonal differences in the diet of giraffe, deciduous trees, shrubs and vines being dominant in the green season, evergreen plants (partly in riverine forest) in the dry season. All the trees and larger shrubs common in the study area were eaten by giraffe, while few records for very small shrubs and none for herbs and grasses were obtained. An analysis of the available vegetation was made in part of the dry-season habitat, and for twenty species the frequency in the habitat was compared with the frequency in the giraffe's diet, revealing selection for or against certain species. Giraffe utilized the upper vegetation layers, where available, but overall c. 50% of all browsing was below 2 m above ground, i.e. within reach of smaller browsers. Results of this study are compared with what is known on food habits of giraffe in other areas. Possible competition of giraffe with other browsers and the relationship between giraffe and their habitat are discussed. Continued survival of giraffe, and other browsers, in Tsavo National Park depends primarily on (1) adequate control of fire, and (2) the impact of future vegetation changes on the amount and variety of available browse plants.     

Impacts of a mineral lick‐centred land use system on woody vegetation cover in an East African Savannah

Evaluation of woody vegetation changes with distance from a salt crater was conducted in the semi‐arid rangelands of southern Ethiopia. Data on live woody plants were collected over three seasons at 0, 1, 4, 6, 9 and 12 km from the salt crater. The density and diversity of woody plants differed significantly (p < .01) along the distance gradient. Six woody plant families were identified of which Fabaceae and Burseraceae were the dominant families. Acacia drepanolobium, Acacia nilotica, Commiphora africana and Acacia mellifera were among the severely encroaching woody species. There were high proportions of seedlings and saplings recorded closer to the salt crater showing a vigorous recruitment by woody plants. Woody plant encroachment along the 12‐km transect ranged from a low to severe encroachment, which could be translated into poor rangeland condition. Changes in soil characteristics increased grazing pressure and sedentary settlement around the salt crater, and the breakdown of traditional institutions seems to be major contributing factors to these vegetation changes. We suggest that severely encroached areas could be improved through a combination of methods such as bush clearing, prescribed fire, browsing animals and proper grazing management.