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.     

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.     

Encroachment of woody plants and its impact on pastoral livestock production in the Borana lowlands, southern Oromia, Ethiopia

Encroachment of woody plants has been among the major threats to the livelihoods of Borana pastoralists and their ecosystem. An approach that integrated vegetation survey and pastoralists’ perception was followed to study the impacts of encroachment of woody plants in the Borana lowlands, Ethiopia. Density of woody species was determined in 192 plots of 500 m². Canopy cover of woody plants was estimated in 123 quadrates of 400 m². Pastoralists’ perception was assessed through group discussions and a semi‐structured questionnaire. Results showed that plant density was 3014 woody plants ha^?1. Cover of woody plants was 52%, indicating an increasing trend from ≤40% cover reported in the early 1990s. It was concluded that the increase of woody plants density and cover has crossed the critical threshold and has entered into the encroached condition. Principal components analysis (PCA) and redundancy analysis (RDA) also showed that woody plants were negatively correlated with herbaceous biomass. Commiphora africana, Acacia melliphera, A. drepanolobium, A. brevispica and Lannea rivae were among the dominant encroachers. RDA revealed that soil nutrients were positively correlated with woody plants density and cover. The pastoralists perceived that encroachment of woody plants had decreased the production of their grazingland. A ban on fire was perceived as the major factor that caused encroachment of woody plants. Re‐utilization of fire and strengthening of traditional rangeland management strategies are recommended.     

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.     

Feeding patterns of elephants at the Atherstone Collaborative Nature Reserve

The African savannah elephant (Loxodonta africana Blumenbach) is a generalist herbivore that relies on widely distributed resources. Vegetation decline, aggravated by these elephants, can compromise local conservation efforts. Thus it imperative to understand the factors that drive them to consume specific plant species and plant parts. The objective of our study was to investigate the feeding patterns of African savannah elephants in the enclosed bushveld savannah at the Atherstone Collaborative Nature Reserve in South Africa. For 1 year, we examined elephant selection of woody versus herbaceous vegetation, and identified which plant species and parts were preferentially consumed. We accomplished this by directly observing feeding elephants, and by utilizing data collected on elephant footprints, dung piles, stripped bark and broken branches. We further conducted vegetation surveys to determine selection frequency relative to species abundance. Elephants showed a preference for different plant parts consumption in the feeding plots. In total, leaves, branches and bark contributed mostly to their diet. Seasonal selection patterns showed an increasing proportion of bark and branch consumption during the dry season.     

Exclusion of livestock grazing and wood collection in dryland savannah: an effect on long‐term vegetation succession

Sahelian savannah faces increasing pressure from human activities, leading to its degradation. The aim of this study was to investigate the possibility of restoration of dryland savannah vegetation by the elimination of disturbance factors on the ecosystem. Is degraded dryland savannah vegetation able to be restored by means of natural succession? What is the timescale for its recovery? The study took place in the Bandia Reserve, 65 km south‐east of Dakar (Senegal), a unique site with two successional stages due to the elimination of uncontrolled exploitation. The vegetation structure of 15 years (15YRS) and 5 years (5YRS) after fencing was compared with vegetation exposed to continuous livestock grazing and wood collection outside the fenced area. Calculated by redundancy analysis, a significant effect of selected areas on the cover of all species was revealed and successional stage explained more than 45% of data variability. Perennial forbs, annual forbs and perennial grasses achieved the highest cover in 5YRS, woody species in 15YRS and annual grasses in the area outside of the fenced reserve. The dominant woody species Acacia seyal, A. ataxacantha, A. nilotica subsp. adstringens and Balanites aegyptiaca reconstituted the dense formation of Acacia bushland by means of natural succession in the 15YRS area.     

Selective disturbance by elephants (<Emphasis Type="Italic">Loxodonta africana</Emphasis>) on eight tree species in a West African savannah

African elephants (Loxodonta africana) are ecosystem engineers in African savannahs, but their role in the modifications of the populations of trees by means of their browsing activities has been poorly studied so far in West Africa. We studied the disturbance of elephants to eight selected species of trees in the Nazinga Game Ranch (Burkina Faso), in 54 transects at the end of dry season 2008. We fitted simple models describing the number of dead individuals for each tree species in relation to their initial population density, and in relation to dung-pile density, in the assumption that there should be a positive relationship between elephant density and dung-pile density. Generalized Linear Model analyses and regression analyses showed that the more dense the woodland cover, the least the percentage of damaged plants by elephants. For each plant species, the initial density and density of elephant dung-piles explained a high proportion of the variance in the density of dead individuals. Stochastic models, generated by a purposely created simple computer program written in GW-BASIC programming language, predicted changes in tree and shrub abundance under different assumptions about elephant numbers. The models suggested that elephant browsing may cause considerable change in the selected plant populations, especially with regard to such species as Acacia gourmaensis, Vitellaria paradoxa and Maytenus senegalensis. These changes may possibly increase the meat harvest from controlled hunting activities, thus improving the income for surrounding communities.     

A case study of apparent conflict between molecular phylogenies: the interrelationships of African elephants

Recent molecular phylogenies of the African elephants suggest that there is an evolutionary structure within Loxodonta africana. Some nuclear results ( Roca et al., 2001 ) support the separation of the forest African elephant subspecies L. a. cyclotis as a species distinct from the savannah elephant L. a. africana, on the basis of the recognition of both forming highly divergent (reciprocally monophyletic) clades. Conversely, a mitochondrial survey ( Eggert et al., 2002 ), while admitting a geographic partitioning of the genetic structure within African elephants, suggests retaining the status quo. They recognize three diagnosible entities (western, central and south‐eastern Africa) with non‐overlapping ranges within L. africana sensu lato. In order to address these conflicting views (historical fragmentation and speciation or isolation by distance, respectively), we have sequenced two datasets of 1961 bp (for 50 elephants) and about 3700 bp, respectively (for 20 elephants) of the mitochondrial DNA for both forms of elephants (cyclotis and africana). They span the cytochrome b gene, the control region and several RNAs. When compared with former mtDNA data, they provide the most comprehensive view of the African elephant phylogeny (78 mtDNA haplotypes, of which 44 are new) and provide the first insight into populations from the Democratic Republic of Congo. The genetic diversity of mtDNA was appraised and the stability of alternative phylogenetic trees was investigated. Our results are inconsistent with both those prior studies. They revealed two highly divergent molecular clades referred to as F and S, that do not conform to the morphological delineations of cyclotis and africana. A non‐negligible proportion of specimens of L. a. africana display haplotypes prevailing in forest elephant populations (clade F). The geographic distribution of clades and areas of their co‐occurrence support the hypothesis of incomplete isolation between forest and savannah African elephant populations, followed by recurrent interbreeding between the two forms. We state that the conclusions of prior studies resulted from insufficient character and/or geographic sampling. We conclude that there is no satisfying argument which can recognize two or more species of African elephants. We briefly comment on the meaning of such an attitude in a conservation viewpoint. © The Willi Hennig Society 2005.     

Woodland changes in Ruaha National Park (Tanzania) between 1976 and 1982

Tree surveys made in 1976 and 1977 were repeated in 1982. Three species were measured: Commiphora ugogensis, Acacia albida, and Adansonia digitata. All three showed significant decreases due to elephant browsing. These results confirm the predictions of the earlier surveys: elephants are causing a rapid and severe change in the rift valley woodlands of the Park.     

The historical development of zoo elephant survivorship

In the discussion about zoo elephant husbandry, the report of Clubb et al. (2008, Science 322: 1649) that zoo elephants had a “compromised survivorship” compared to certain non-zoo populations is a grave argument, and was possibly one of the triggers of a large variety of investigations into zoo elephant welfare, and changes in zoo elephant management. A side observation of that report was that whereas survivorship in African elephants (Loxodonta africana) improved since 1960, this was not the case in Asian elephants (Elephas maximus). We used historical data (based on the Species360 database) to revisit this aspect, including recent developments since 2008. Assessing the North American and European populations from 1910 until today, there were significant improvements of adult (≥10 years) survivorship in both species. For the period from 1960 until today, survivorship improvement was significant for African elephants and close to a significant improvement in Asian elephants; Asian elephants generally had a higher survivorship than Africans. Juvenile (<10 years) survivorship did not change significantly since 1960 and was higher in African elephants, most likely due to the effect of elephant herpes virus on Asian elephants. Current zoo elephant survivorship is higher than some, and lower than some other non-zoo populations. We discuss that in our view, the shape of the survivorship curve, and its change over time, are more relevant than comparisons with specific populations. Zoo elephant survivorship should be monitored continuously, and the expectation of a continuous trend towards improvement should be met.     

Effects of gut treatment on recovery and germinability of bovine and ovine ingested seeds of four woody species from the Sudanian savanna in West Africa

Passage rate through the digestive tracts of zebu cattle and sheep, and subsequent germination of egested seeds of four woody species from the Sudanian savanna, Acacia dudgeoni, Acacia seyal, Burkea africana and Prosopis africana, were studied. The result indicates large differences in passage rate among woody species, as well as between animals. The values ranged from 46% to 87% for seeds ingested by cattle while the lowest passage rate was 2.3% and the highest being 74% for seeds ingested by sheep. Among plant species, seeds of Prosopis africana had the highest passage rate through the digestive tract of both cattle and sheep. Seed passage through the gut showed a significant positive correlation with seed mass and thickness for cattle and sheep, respectively. The gut treatment and the retention time in the gut did not improve germination capacity and the speed of germination of dormant seeds. For non-dormant seeds of Acacia dudgeoni, the germination capacity was higher for seeds ingested by cattle than sheep. The speed of germination was also significantly higher for egested seeds than the control. It can be concluded that large herbivores could play an essential role in long distance dispersal of seeds. Gut treatment alone was not effective in breaking seed coat-imposed dormancy, although it enhanced the rate of germination of non-dormant seeds. To get a complete picture of the effect of frugivore on the release of seed dormancy, the combined effect of initial mastication and subsequent gut treatment needs to be investigated.     

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.)     

How does a re-colonizing population of Asian elephants affect the forest habitat?

The Asian elephant Elephas maximus is currently re-colonizing the Bardia National Park in lowland Nepal. We studied their impact on woody vegetation in the nutrient-rich floodplain and in the relatively nutrient-poor sal forest. The types and extent of tree impact were recorded along fixed-width transects (335 km). Species composition, density and size classes ≥8 cm diameter breast height (dbh) were recorded in 15-m radius random plots ( n =95). Impact was higher in the floodplain complex than in the sal-dominated forest. Our hypothesis that elephants were more selective on species in the nutrient-poor sal forest was only partly supported; the niche breadth of impacted trees was slightly higher in the floodplain complex. Pushed-over trees accounted for the highest proportion of impact (55%), followed by killed trees (39%). Of the pushed trees, 10% were not used for food. Among food trees, elephants selectively impacted size class 12–16 cm dbh, whereas non-food trees were impacted independently of size. A large proportion of the freshly browsed trees had been felled previously, indicating that most felled trees survived, enabling elephants to feed on them again. This may reflect an evolutionary adaptation among long-lived species with high site fidelity. Owing to preferential use but low abundance, two species in sal forest, Grewia spp. and Desmodium oojeinense , were found to be particularly vulnerable to local extinction due to elephants. Although the elephants had impacted a large number of species (62, 73% of all), 56.4% of the impacted trees consisted of Mallotus phillippinensis . A recently observed increase in the density of M. phillippinensis and the concurrent reduction of the hardly utilized Shorea robusta indicates that the rapidly growing elephant population may modify the composition of the forest by increasing its preferred food species.     

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.     

Human settlement drives African elephant (Loxodonta africana) movement in the Sebungwe Region,Zimbabwe

Understanding the key drivers that influence the potential distribution of herbivore species in changing landscapes has been at the centre of enquiry in wildlife science for many decades. This knowledge is particularly important for keystone species like the African elephant (Loxodonta africana) whose population is declining even in conservation areas. The Sebungwe Region is part of the Kavango‐Zambezi Transfrontier Conservation Area and supports ~4,000 elephants. The Sebungwe Region has lost an estimated 76% of its elephant population over the years. This study aimed to determine how the elephant distribution in the Sebungwe Region was affected by human settlement and whether the patches for elephant distribution were large enough for elephant habitation. The prediction of the potential distribution of the elephant was based on presence‐only data modelled through an ensemble algorithm that combined several candidate models to enhance predictive ability. We observed that human settlement drives the potential distribution of elephants in the Sebungwe Region (test AUC = 0.95), and patches from the model were on average <1.5 km². Our results provide initial insights into the key habitat factors that drive distribution of elephants in the Sebungwe landscape. Future conservation of the elephant could benefit from our study through systematic planning of settlements, which might help minimise human interaction with wildlife.     

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.     

Diversity and abundance of insect visitors to flowers of trees and shrubs in a South African savannah

This study presents the results of a landscape‐scale survey for insect floral visitors in the Skukuza Ranger District, Kruger National Park, South Africa. Floral visitors were sampled from flowering trees and shrubs along linear transects spanning the entire district. Six plant species were sampled in the late dry season (Acacia grandicornuta Gerstner, A. nigrescens Oliver, Cassia abbreviata Oliver, Combretum hereroense Schinz, Combretum zeyheri Sonder, Euclea divonorum Hiern), and eleven plant species were sampled during the rainy season (Acacia exuvialis Verdcourt, A. grandicornuta Gerstner, A. nilotica (L.) Willdenow, A. tortilis (Forsskal) Hayne, Dichrostachys cinerea Miquel, Flueggea virosa (Roxburgh) Baillon, Grewia bicolor Jussieu, G. flava De Candolle, G. flavescens Jussieu, G. monticola Sonder, and Peltophorum africanum Sonder). Coleoptera, Hymenoptera and Lepidoptera comprised the majority of floral visitors, while species of Blattodea, Diptera, Hemiptera and Neuroptera also occurred on flowers. Known or likely pollinators include bees (Hymenoptera: Apidae, Halictidae and Megachilidae) and scarab beetles (Coleoptera: Scarabaeidae). These plant species appear to have generalist pollination systems, with the exception of species of Grewia L., which appear to be pollinated primarily by bees. A provisional plant–pollinator food web is presented for the eleven species of trees and shrubs which flower during the rainy season.     

Characterization of tetranucleotide microsatellite loci in the African Savannah Elephant (Loxodonta africana africana)

Most African elephant (Loxodonta africana africana) populations are isolated and thus threatened by a loss of genetic diversity. As a consequence, genetic analysis of African elephant populations will play an increasing role in their conservation, and microsatellite loci will be an important tool in these analyses. Previously published sets of polymorphic microsatellites developed for African elephants are all dinucleotide repeats, which are prone to typing error. Here, we characterize 11 tetranucleotide microsatellite loci in the African elephant. All loci were polymorphic in 32 faecal samples and two tissue samples from 33 individual African savannah elephants.     

The hard lives of trees in African savanna—Even without elephants

The ongoing loss of large trees and densification of shrubs are two prevalent processes that take place in African savannas, with profound consequences for their structure and function. We evaluated herbivore impacts on savanna woody communities using a long-term exclosure experiment in the Kruger National Park, South Africa, with three treatments: the exclusion of large mammals only (i.e. elephant and giraffe), exclusion of all herbivores larger than a hare, and areas open to all herbivores. We asked three questions: (1) How did variable exclusion of herbivores affect woody density and structure across the catena (i.e. riparian, sodic and crest vegetation)? (2) Did the exclusion of herbivores result in unique woody species composition? (3) Did herbivore exclusion result in a higher proportion of palatable species? After 17 years, we found that herbivores mainly affected the heights and densities of existing species, rather than leading to turnover of woody species assemblages. Although densities of individuals increased in the full exclosure (350 ha⁻¹), the change was more moderate than expected. By contrast, mixed mega-and meso-herbivores decreased the number of trees and shrubs (decreases of 780 ha⁻¹) via a variety of physical impacts. Meso-herbivores alone, on the other hand, had less impact on individual density (i.e. no change), but limited average height growth and canopy dimensions in certain habitat types. Where elephants are present, they are effective at reducing the density of woody stems to the point of counteracting woody encroachment, but at the same time are actively preventing the persistence of large trees (>5 m) as well as preventing trees from recruiting to larger size classes. However, the lack of massive recruitment and woody cover increases with elephant exclusion, especially for more preferred species, suggests that factors beyond elephants, such as dispersal limitation, seed predation, and drought, are also acting upon species.     

Responses of woody vegetation to exclusion of large herbivores in semi‐arid savannas

The Nkuhlu large‐scale long‐term exclusion experiment in Kruger National Park was designed to study the long‐term effects of large herbivores on vegetation. One treatment excludes elephants, another excludes all herbivores larger than hares and another one comprises an open, control area. Vegetation monitoring was implemented in 2002 when a baseline survey was conducted prior to exclusion. Monitoring was repeated 5 years after exclusion. Data from the surveys were analysed to establish how structure and composition of woody vegetation had changed 5 years after herbivore exclusion. The analysis showed that neither plant assemblage nor mean vegetation height had changed significantly since exclusion. However, both species richness and density of woody plants increased 5 years after exclusion of all large herbivores, but not after the exclusion of elephants alone. One already common species, Dichrostachys cinerea, became more common after excluding all large herbivores compared with either no exclusion or elephant exclusion, possibly leading to competitive suppression of other species. Species other than D. cinerea tended to either increase or decrease in density, but the changes were insufficient to induce significant shifts in the overall assemblage of woody plants. The results indicate that after 5 years of exclusion, the combined assemblage of large herbivores, and not elephants alone, could induce changes in species richness and abundances of woody plants, but the effect was so far insufficient to induce measureable shifts in the assemblages of woody plants. It is possible that assemblages will change with time and increasing elephant numbers may amplify future changes.