Most elephants in South Africa live in enclosed areas such as the Tembe Elephant Park in Maputaland. The Park also protects sand forest. This can create a conflict of interest as elephants may influence species typical of these forests. To assess the effects that elephants may have for vegetation, I compare variables of similar plant communities inside and outside the Park. I then compared the space and landscape utilization of elephants living in the Park with those of free-ranging elephants living in southern Mozambique. In the final analyses, I used meta-analytical methods to interpret my findings. Woody seedlings showed no measurable response to tree canopies that elephants have altered, but the response of grasses and woody saplings depended on the landscape type. In closed woodlands, elephants generated gaps in the canopy layer that increased structural heterogeneity. These gaps favoured the establishment of grasses, and along with herbivory, may have been responsible for reduced occurrence of woody saplings. In the open woodlands, elephants and frequent hot fires in the Park apparently homogenised this landscape. In this case, altered tree canopies reduced grass and woody sapling presence. The species compositions of sand forests, closed woodlands and open woodlands between inside and outside the Park differed. However, tree and shrub densities, their abundance-incidence and rank-abundance relationships were similar for a given landscape inside and outside the Park. Ecological events operating at larger scales, such as seed dispersal and droughts, mask the influence elephant have for these community variables. Elephants in the Park had smaller home ranges than free-ranging elephants living in southern Mozambique. The size of these home ranges were however, similar to that predicted by rainfall, as suggested by my analysis of data collected across southern Africa. The elephants that roamed freely in southern Mozambique prefer closed woodlands throughout the year. However, elephants confined to the Park avoided reed beds (with natural surface water) in the dry season and showed no landscape preference in the wet season. My meta-analysis on the effects of elephants on other taxa included 230 peer-reviewed studies. These were published over a 40-year period and included information from 74 sites. From only those studies used in the effect size calculations, when conducted over a period of less than 5 years show a negative impact while those conducted over longer periods show a neutral effect. Site-specific differences, such as rainfall, may also influence the effect elephants have for plants. Twenty of the 230 studies shared more than 50% of all citings. The majority (16 of the 20) claimed that elephants had a negative influence for plants. This is in contrast with the findings of all studies included in the analysis – half of these concluded a positive effect and the other half a negative effect. In short, elephants do not decrease the diversity of other species present in the system, despite their adverse effects for individual trees. Elephants affect ecosystems at small scales. Providing an opportunity for elephants presently living in Tembe Elephant Park to disperse across their former ranges may negate negative influences on sensitive vegetation in the Park. / Thesis (PhD (Zoology))--University of Pretoria, 2007. / Zoology and Entomology / unrestricted

In Central Africa, wildlife populations are increasingly influenced by humans, even in protected areas. This raises the question how spatial patterns of wildlife abundance are affected by human activities and habitat and how these patterns change over time. I address these questions by developing spatial models combined with line transect survey data in two forest sites in Central Africa. In the Odzala National Park in the Republic of Congo, I examine elephant dung abundance data in relation to human threats and protection. In the Okapi Faunal Reserve in the Democratic Republic of Congo (DRC), I developed spatio-temporal models for elephants and other forest ungulates to examine temporal changes in their densities as a result of changes in human impact in the context of a civil war that took place in the region between the two surveys. Covariates related to human influences dominated the observed patterns at both sites. In Odzala, elephant dung densities were mainly influenced by protection. They were higher inside the protected area and closer to anti-poaching patrol routes. In the Okapi Faunal Reserve, populations of all ungulate species declined severely between the two survey periods. Declines in elephant abundance were relatively higher closer to the park boundary and areas of intense human activity. After the war, elephant densities were higher in a small area in the centre of the park that may have acted as a refuge. Forest duikers also declined, but the spatial pattern of their decline was different than that of elephants. Densities dropped more in the southern part of the reserve, probably due to pre-exisisting higher levels of hunting there. Besides explaining spatial patterns of abundance, spatial modeling was shown to be useful in improving the precision of density estimates and in predicting densities across a surface in the Odzala National Park. In summary, humans overwhelmingly determined the distribution and abundance of ungulates in both sites. The civil war in DRC led to a dramatic increase in elephant poaching for ivory which caused a major decline in elephant populations. It aggravated the bushmeat hunting of duikers whose populations also declined sharply. / Science, Faculty of / Zoology, Department of / Graduate

When elephants leave primary protected areas (PPAs), such as national parks and game reserves, they may come into conflict with people residing on the adjoining land. In this study, I attempted to determine why African savannah elephants leave the PPAs in which they were collared. To accomplish this, I used telemetry locations of collared elephants in PPAs throughout southern Africa and investigated whether a range of intrinsic and extrinsic variables could explain why elephants crossed the boundaries of the PPAs. Adjoining many of the PPAs were secondary protected areas (SPAs), which consisted of community conservancies, and collectively with the PPAs formed clusters of protected areas. Most (45 of 49) elephants roamed beyond the PPAs but they remained within the clusters of protected areas. The elephants utilised both the PPAs and the SPAs and appeared to not feel threatened when using the SPAs. The reasons for elephants leaving PPAs varied both seasonally and between the sexes. The females roamed beyond the PPAs more during the wet season than the dry season, whereas, for males there was no seasonal difference. During the wet season, female and male habitat selection was similar within and beyond the PPAs. During the dry season, more females and males beyond the PPAs selected for areas close to people, which could be indicative of water. The proportion of male and female home ranges beyond PPAs did not increase with increasing density of elephant populations within the PPAs, nor did the proportion of female home ranges beyond PPAs increase with increasing population growth rate of elephant populations within the PPAs. Therefore, high numbers of elephants within the PPAs did not drive elephants beyond the boundaries. Contrastingly, the proportion of male home ranges beyond the PPAs did increase with increasing population growth rate. However, the results were inconclusive due to small sample size. This study confirms that elephants are utilising the SPAs as well as the PPAs. Therefore, the importance of including the SPAs in conservation actions for elephants cannot be over-emphasized. / Dissertation (MSc)--University of Pretoria, 2013. / Zoology and Entomology / unrestricted

Please read the abstract in the 00 front of this document. / Thesis (PhD)--University of Pretoria, 2009. / Veterinary Tropical Diseases / unrestricted

Elephants can cause people harm and destroy property in communities livingclose to national parks. Having an automated system that can detect and warnthe people of these communities is of utmost importance in order for human andelephant coexistence. Elephants heavy profile and damped footsteps induce lowfrequency ground waves that can be picked up by geophones. In the thesis twomain problems are investigated, detecting if the geophone measurements contain an elephant footstep and calculating the direction of the elephant footstep.A real time system is built containing a sensor array of three geophones. By analyzing the frequency content of the geophone measurements, elephant footstepscould be detected. The system in capable of detecting elephants situated up to40 meters away from the geophones. Utilizing the sensor array, a direction to theelephant was estimated using triangulation. Two methods of triangulation wereinvestigated. At 15 meters away, the estimation deviated with only a few degrees.At 40 meters away, the estimation was at least good and consistent enough to geta general idea of where the elephant was coming from.

Although the unprotected lands of northern Tanzania support large numbers of elephants, and provide critical linkages for wildlife movements across the region, there is little information on the dispersal patterns of elephants in these unprotected lands. Our home range measures (100% MCP) of 21 elephants with satellite collars in four study regions were highly variable (191 to 3,698 km2). Home range sizes (95% fixed kernel) of bulls were typically larger than those of females, and wet season ranges were typically larger than dry season ranges. There were large differences in average home range sizes reflected varying strategies for obtaining food and water and avoiding humans. All eight radio-collared elephants (3 bulls, 5 females) in the West Kilimanjaro study region crossed the Tanzania-Kenya border, but typically elephants crossed more frequently in the wet than the dry season, and bulls crossed 47% more frequently than females. These extensive transboundary movements indicate that the elephant populations of West Kilimanjaro and Amboseli NP constitute a single transboundary population. Based upon 14,287 fixes from eight collared elephants, the vast majority of time was spent in unprotected ( x=91.5%) versus protected (x =8.5%) areas. Amboseli NP was visited by all eight elephants and was the protected area most utilized ( x=8%, range 2-24%). Based upon the movements of 15 GPS-collared elephants in northern Tanzania, we identified eight areas that we considered important for wildlife conservation corridors/linkages for elephants. Our conservation priorities for these corridors were based upon the levels of threats and conservation potential. Community interviews and hilltop surveys were used in two Maasai villages to determine the extent of wildlife conflict, community attitudes towards elephants, and if elephants were using a vegetation corridor to move between Tanzania and southern Kenya. Elephants were the most problematic wildlife species and were considered a nuisance. However, they believed they attracted tourists, and generally did not believe elephant numbers should be reduced. Based upon elephant conflict and use and the communities' need to maintain areas for cattle grazing and medicinal plant collection, the two communities established the first wildlife conservation corridor in Tanzania.

Land use changes have been shown to have significant effects on wildlife species. Sixty three percent of the national Parks and Reserves in Kenya are located in the savannas. Because of the seasonality associated with savanna ecosystems, 75% of the wildlife species and numbers in Kenya occur in savanna nonpark areas. Therefore, conservation of wildlife in Kenya has to address the changes that are taking place in savanna areas outside parks. I studied land use changes and their effects on elephant habitat quality in Amboseli Basin, Kenya. I used visual interpretation to analyze land use changes from satellite images for 1975, 1988, and 1993. I determined that during the evaluation period, conversion of areas to agricultural land has been unidirectional. The hectarage under cultivation was 2,937, 10,950, and 24,476 for 1975, 1988, and 1993 respectively. Trend analysis seems to suggest that during the evaluation period, conversion of areas to agricultural land has followed an exponential function (R² = 0.99) in Amboseli Basin. The area under cultivation was 6.9% of the total area studied. This is small but significant considering that agricultural land was almost exclusively located in areas that form the dry season fall back areas. Such areas are important for the survival of elephants and other species during critical periods. I developed a dry season habitat suitability index (HSI) model for the African elephant based on the density of acacia trees (# of trees > 5 cm dbh/ha) and distance (km) to natural sources of water in the basin. The amount of good quality habitat (i.e., HSI > 0.6) declined from 74,666 ha in 1975 to 54,890 ha in 1988, to 23,208 ha in 1993. This is a drop of 51,890 ha (65.5%) of good quality habitat in the basin. On the other hand, low quality habitat (HSI < 0.2) increased by 272% between 1975 and 1993. The weighted HSI values in the basin showed a decline, as did the habitat units for the 3 evaluation years. The weighted HSI declined by 0.13 between 1975 and 1993, while the total habitat units (ha) declined by 40,567 ha during the 18 year period. It appears that elephant habitat quality has steadily declined in the Amboseli Basin during the period considered in this study. The use of nonpark areas by elephants leads to direct interaction and conflicts with humans. Between June 1996 and July 1997, I recorded a total of 489 elephant damage incidents while the Amboseli National Park authorities recorded 143. The damage types were crops, livestock deaths, and human deaths and injuries. The majority of the damage cases involved crop depredation. The Amboseli National Park authorities significantly under-reported the number of elephant damage incidents in the basin (P < 0.0001). / Ph. D.

Animals are restricted in their diets by several factors, most notably the availability and quality of resources. Variation in resource availability causes herbivores to shift their diets seasonally and spatially. Elephants (Loxodonta africana), are known to have extensive impacts on plant communities, altering ecosystem functioning and causing a decline in biodiversity. In enclosed areas, these impacts are increased leading to a decline in resource availability and presumably resource quality. In the Addo Elephant National Park, the Main Camp section has a history of high elephant impacts and therefore reduced resource availability. Whereas, the recently added Colchester section has greater resource availability, due to the absence of elephants in this section since the fencing of the Park. This study investigated the changes in diet (diet breadth, preference and diet quality) of elephants due to an increase in resource availability. Three alternative hypotheses were contrasted: 1) elephants as generalist foragers, 2) elephants as optimal foragers, or 3) elephants learning foraging behaviour. Using microhistological analysis, the diets of elephants were described over five sampling periods (August 2010 – February 2014) in both sections. Forage availability was estimated using a modified line-intercept method, and was used to determine changes in preference by relating forage availability to use. In the Colchester section the diet breadth of elephants increased, and was coupled with a high initial variation between the diets of elephants, which decreased in subsequent sampling periods. This supported the elephants learning foraging behaviour hypothesis. However, there was no increase in diet preference by elephants in the Colchester section, which supported the elephants as generalist foragers hypothesis. There was also no difference in the diet quality of elephants in the Main Camp and Colchester sections, which did not support any of the three hypotheses. The elephants learning foraging behaviour hypothesis is proposed to be the link between the alternate two hypotheses, and given enough time, either of the two could be supported. The lack of difference in preference and diet quality between elephants in the Main Camp and Colchester sections is hypothesised to be due to the population level (not measured for individuals) at which these were measured. Microhistological analysis of faeces was used to describe the diet of elephants, which was compared to the diet described by DNA metabarcoding. Microhistological analysis is a traditional, favoured technique used in describing the diet of wild herbivores, whereas DNA metabarcoding is a relatively new and untested technique. These two techniques have not yet been compared in the diet of megaherbivores. Results indicated that microhistological analysis identified significantly more grass in the diet of elephants, than DNA metabarcoding did, which was expected as previous studies also found overestimation of grasses. Microhistological analysis identified more plant families in the diet of elephants, than DNA metabarcoding. Most of the differences between the two techniques can be attributed to the difference in taxonomic resolution, which was due to the lack of a complete reference collection for DNA metabarcoding. Although either of the two techniques can be used to describe the diet of elephants, the most reliable results would be obtained when using both techniques. The findings of this study suggest that due to the high initial variation between the diets of elephants, with an increase in resource availability, the impacts will also initially be highly varied. This suggests that identifying plant species to monitor elephants impacts initially will be difficult. However, important plant species, or those known to be vulnerable to elephants impacts should be carefully monitored initially and monitoring should not only occur annually, but also seasonally.

Elephants, as megaherbivores and keystone species have major impacts, both positive and negative, on ecosystems such as the Subtropical Thicket of the Eastern Cape. The feeding behaviour of elephants was quantified so as to firstly determine the preferred feeding heights of elephants, and secondly to determine if feeding behaviour and impact varies with size and sex. The preferred feeding height was determined with experimental feeding trials. Feeding height, pluck size, foraging rate and the proportion of plant material discarded were used to test for differences between elephants of different sizes and sexes in the Addo Elephant National Park. The influence of plant growth form on sex-specific feeding was also considered. Elephants preferred to feed at the lowest heights. The preferred feeding height was not related to body size. A wide range of feeding heights was recorded including and extending beyond, both the preferred and maximum feeding height of co-existing indigenous browsers. There was no difference in feeding height between the sizes and sexes. In free ranging conditions, all elephants fed at levels above the preferred foraging height when browsing. There was no difference in feeding behaviour between the different sized elephants, or between males and females. Adult elephants however exhibited larger ranges of feeding heights, pluck sizes, foraging rates and intake rates, including the maximum and minimum values. Growth form influenced the feeding of male and female elephants differently. Elephant feeding behaviour appears to limit opportunities for resource partitioning by way of feeding height segregation. Elephants are also capable of dominating the browse resources available to co-existing browsers by removing large amounts of forage at lower heights. The extent of elephant impact seems to be comparable between sizes and sexes, although adults and especially larger males are often able to use foraging opportunities that other elephants can not effectively utilize. Findings suggest that the feeding heights of elephants are determined by forage availability and reflect the diet in terms of browse or grass. The findings can be used, together with browse inventory methods, to determine browse resources available to elephants, and can also be used to develop monitoring tools to assess the impacts of elephants on plants.

The development of small (<300 km²), private game reserves has become a trend, not only in the Eastern Cape Province, but also elsewhere in South Africa as a result of a shift in land use practise from agriculture to ecotourism. The resultant re-introduction of elephants to many of these reserves has lead to management concerns because of the limited research on small reserves regarding their impact on the vegetation. In this study I assessed the space use, habitat selection, diet and impact of two elephant populations on the vegetation in the Eastern Cape Province between February 2004 and March 2005. Home range sizes were calculated using the kernel utilization distribution method. Home range sizes for elephants on Kwandwe were significantly larger during summer than winter (p<0.05). There was no significant difference between the home range sizes of the herds and males within a season and during summer the elephants utilized about 75 % of the reserve and only 54 % during winter. On Shamwari, the herd utilized about 92 % of the reserve during summer and 83 % during winter; while the males utilized 76 % of the reserve. Core areas for both elephant populations shrank from summer to winter and were concentrated around the permanent water sources on each reserve. Habitat selection was assessed using χ² tests and Bonferroni confidence intervals. On Kwandwe, there was a significant difference between observed and expected use of vegetation types (p<0.05) and karroid shrubland was strongly avoided by both herds during summer and winter. The preferred vegetation types of the males ranged from relatively open (short euphorbia thicket, bushclump karroid thicket and karroid shrubland) to completely open (old lands). On Shamwari, subtropical thicket, bontveld and montane grassland were avoided; while primary and secondary acacia thicket, riverine thicket and cultivated lands were preferred. The predominant vegetation type in the home ranges of herds on Kwandwe and Shamwari was subtropical thicket. The diet was assessed by direct observations over two seasons and dietary preferences were calculated. There was a significant difference in the frequency of occurrence of plants in the diet on the two reserves (p<0.05) and no significant effect of time of day or season (p>0.05). Seventeen woody plant species were utilized on Kwandwe and 23 species were utilized on Shamwari. Grass constituted a significantly greater percentage of the diet in summer than winter (p<0.05). Elephants on Kwandwe showed a selective preference for Ozoroa mucronata, Pappea capensis and Acacia karroo; while on Shamwari, A. karroo was selected. Transects were conducted in two different vegetation types on each reserve so as to assess the impact of elephant on the vegetation and damage scores were then calculated from these data. There was no significant effect of vegetation type or elephant density on mean damage scores in Kwandwe (p>0.05). Five hundred and seventy-eight plants were assessed in the subtropical thicket vegetation type and 225 plants were assessed in the savanna-type vegetation, with more than half the trees showing low levels of damage that could not only be attributed to elephants. Mean damage was highest for Portulacaria afra and Pappea capensis in subtropical thicket and for Rhus spp. in the savanna-type vegetation. On Shamwari, 408 plants were assessed in subtropical thicket and 215 in the savanna-type vegetation, with more than 70 % of trees showing low levels of damage. There was a significant effect of plant species and elephant density on the mean damage scores in subtropical thicket, with Aloe ferox showing more damage than the other plant species (p<0.01). In the savanna-type vegetation, A. karroo was the most severely damaged. Overall, damage was greater in the thicket vegetation type compared to the more open vegetation type on both reserves.