Elephants' complex societies, well-developed communication systems, evolutionary history and close working relationship with humans make them an important species for studies of cognition but research on elephant cognition is sparse. In this thesis I aim to illuminate the social cognition involved in the interpretation and production of visual signals by African elephants (Loxodonta africana). My results are intended to contribute to the cross-species literature on social cognition and help to elucidate wild elephant social behaviour. I studied captive elephants, housed at an elephant-back safari company in Victoria Falls, and wild elephants in Hwange National Park, both in Zimbabwe. Wild elephants display a vast array of postures, actions and signals. I found that elephants recognise visual attentiveness in others when they signal silently, producing more signals when their audience can see them, and using the body and face orientation of an audience to judge their attention. When responding to typically human visual signals, elephants immediately responded correctly to deictic gestures, including variants of pointing that they were unlikely to have already experienced. These results indicate elephants' astonishing sensitivity to even subtle social cues. I found no indication that elephants reason about mental states such as false beliefs, or rationality; however, limitations of the experimental design meant I was unlikely to find such an ability even if it is present in elephants. Furthermore, I discovered that elephants have a form of referential indication in their natural communication in the wild. Elephants match their direction of attention with a type of trunk action produced by a group member. Attending to human-like signals, and interpreting them as communicative is an advantage for any animal working with humans and that ability might explain the choice of species that are ancestors of today's domestic animals.

Captive populations of Asian elephants (Elephas maximus) in North America are not self-sustaining, and increasing reproductive success within captive populations is a high priority. The ability to accurately predict parturition can have a direct impact on elephant welfare. Elephants in captivity often require significant preparation and management throughout the birthing process, and complications during labor and delivery can necessitate immediate intervention, including stillbirth, protracted labor, maternal aggression towards a newborn calf, and dystocia. Being able to predict when parturition will commence can ensure appropriate staff is available and adequate monitoring is performed. Routine endocrine sampling can be used to predict parturition in Asian elephants, with a drop in progesterone (P4) to baseline levels signaling parturition in 2-5 days. However, we determined this method is not without limitations, and it is not used in all institutions that house elephants. As changes in hormones regulate and alter behaviors, we investigated behavioral indicators as an additional management tool for predicting parturition, a time of drastic hormone changes. We conducted a study of five pregnancies in Asian elephants at the Oregon Zoo, U.S.A, and Taronga Zoo, Australia, between 2008 and 2012. In Chapter 2, I evaluated progesterone (P4) and cortisol levels across three time periods: Baseline; Pre, (the week preceding the drop in P4); and Post, (the period after the P4 drop). Levels of P4 were significantly lower, and levels of cortisol were significantly higher in the days just prior to parturition. I found considerable intra- and inter-individual variation in both endocrine profiles, which can make endocrine assessments difficult to interpret in real time. In Chapter 3, I investigated whether behaviors in the preparturition period could be predictive of impending parturition in the Asian elephant. ANOVA results indicated a significant difference in the amount of time that elephants spent walking backwards across three time periods (F(2) = 3.723, p = 0.033), with the behavior increasing as parturition approached. These results were supported by a non-parametric Kruskal- Wallis. Using a generalized linear mixed model (GLMM), I found that as P4 levels decrease, walking backwards behavior significantly increases. In Chapter 4, I evaluated investigative trunk behaviors, or "trunk checks", directed towards the temporal gland near the ear, mammary glands, vulva and anus of the pregnant dam. Investigative behaviors included both self-directed behaviors and those sent from herd mates towards the pregnant dam. Self-directed behaviors are most likely associated with physical changes in the pregnant dam, such as using the trunk to pull on swollen teats. Other-directed behaviors may stem from chemo-sensory signaling or other types of communication between herd mates, such as detecting changes in progesterone or cortisol. I ran GLMM and found that four trunk-check behaviors varied significantly with P4 and/or cortisol profiles. These were: self-checks of mammary glands increased with decreasing P4 levels; herd-mate-checks of mammary glands increased with decreasing P4 levels; self-checks of vulva increased with decreasing P4 levels and increasing cortisol levels; herd-mate-checks of anus increased with increasing cortisol levels. In Chapter 5, I evaluated activity budget behaviors in the pregnant elephants. Generalized comparisons were made to published activity budgets of typical captive Asian elephants. I report that activity budgets are within the range of normal activity, though I note a high level of inter-individual variation. In addition, I compared two sampling techniques, including one-zero and instantaneous sampling, that were used for activity budget data collection. I discuss the different results obtained by each sampling technique. These results are a very promising indication that behaviors, including walking backwards and multiple trunk-check behaviors, are changing over time or with parturition-related hormone profiles. We recommend that keepers, veterinary staff, and other observers that are familiar with the regular behavioral repertoire of a pregnant female should pay close attention to these highlighted behaviors. Keeping track of these behaviors, especially in conjunction with P4 and cortisol tracking, can help staff refine existing windows of expected parturition.

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.

Compromised welfare and wellbeing of elephants (Loxodonta africana and Elephas maximus) in captive facilities are significant and global problems. The period between birth and two years old is crucial for calf survival and social and environmental learning. Behaviour and developmental processes among captive elephant calves in these first years were compared with those seen in wild calves. Wild elephants calves develop within a complex, varied social context and provide one reference for normal patterns of development. Such comparisons enable insights into welfare at captive facilities. Eleven captive elephant calves born at three UK facilities were studied from birth to 18 months (AsianN=6; AfricanN=5). Older calves (AsianN=2; AfricanN=2) were also sampled up to 3.5 years; making a total of 15 calves studied from 2009 to 2014. Due to the small sample size, the 11 younger calves were also discussed as individual case studies. By 2017, only two of these case study calves were both alive and not orphaned. Three additional calves (AsianN=1; AfricanN=2) died on their day of birth and were not sampled. This small sample highlights the ongoing lack of self-sustaining populations of captive elephants. This thesis collated systematic behavioural observations on captive calves across 373 days (483.5hrs). Calf maintenance activities (feeding, resting, moving), associations with mother and others, interactions and calf play were compared with behavioural observations of wild AsianN=101 (74hrs, Uda Walawe, Sri Lanka) and wild AfricanN=130 (252hrs, Amboseli, Kenya) calves from ~birth to five yrs. Mothers’ (captive: AsianN=4; AfricanN=4; wild: AsianN=90; AfricanN=105) activities were also recorded to explore synchrony with calves. Captive calves raised by their mothers had similar activity budgets to those seen in the wild. Expected age-related declines in suckling were found in captivity. However, captive calves were more independent than wild calves for their age in distance from mother and spent significantly more time in play. A Decision Tree for whether to breed elephants in captivity was developed; benefits that a calf potentially brings to companions, e.g. multi-generational matrilineal groups, enabling social bonding and reducing abnormal behaviours, were considered against space required for families to grow and divide naturally over time, as well as ensuring that captive-bred males are socially sustained. It was recommended that facilities invest in future enclosure/housing designs which permit: free-access to other elephants; 24hr trickle feeding; juvenile males allowed to stay with their maternal group for longer, encouraging learning opportunities and further retaining age-structure/composition. Conversely, facilities unwilling to house a male or provide appropriate group size/composition are recommended to cease breeding.

Darwin was troubled by elaborate male traits observed in many species that are seemingly maladaptive for survival, the peacock’s tail being the most iconic of all. He wrote "The sight of a feather in a peacock’s tail, whenever I gaze at it, makes me sick" because it challenged his theory of evolution by natural selection for adaptive traits. The extreme length of the tail may render a peacock more vulnerable to predation and therefore maladaptive for survival. To account for the evolution of apparently maladaptive traits he proposed the theory of sexual selection, wherein, traits that directly enhance mating success may be selected for, either as weapons in male-male competition for mates or as ornaments preferred by females. Male and female elephants in the proboscidean evolutionary radiation have had tusks and show extreme exaggeration in size and form. However, tusk in the Asian elephant (Elephas maximus) is sexually dimorphic as it is expressed only in the males, hinting at a possibility that opposing selection (sexual selection advantage to males and natural selection disadvantage to females) may have been the processes behind this pattern of tusk expression. Intriguingly, tuskless males (male dimorphism with respect to tusk) also occur at fairly high frequencies in some Asian elephant populations (∼50% in norteastern India and ∼95% in Sri Lanka). Theory states that dimorphic males can also occur in a population in stable frequencies as a consequence of sexual selection. I explored sexual selection on elephant tusks as possible mechanism leading to the observed patterns of tusk dimorphism in the elephants. All elephant populations on earth have been harvested for ivory, therefore, artificial selection (selective poaching of tusked elephants for ivory) is another possible cause of tusk dimorphism. I developed mathematical models of population genetics, population dynamics and conducted field observations of mating behavior of Asian elephant in Kaziranga National Park, Assam to understand the evolution of tusk dimorphism in elephants. Darwin’s sexual selection theory was controversial when proposed in 1871 and continues to remain so in 2014. In the introduction of my thesis I have discussed Darwin’s two classical mechanisms of sexual selection, namely, male-male combats for mates and female mate choice based on male traits. The latter was viewed with considerable skepticism by his con-temporary Alfred Russell Wallace and more recently deemed "fundamentally flawed" by Joan Roughgarden. Therefore, I have also discussed the arguments against female mate choice for male traits found in literature. I have reviewed current knowledge about sexual selection for sexually dimorphic male traits of body size and musth, in the African and Asian elephant and state why I have hypothesized that tusks may also be under sexual selection. Sexually selected traits are expected to be genetically determined, therefore, I explored mathematically (Chapter 1) the genetic basis of evolution of sexual dimorphism. Fisher proposed that sexually selected male display traits originate in both the sexes but are suppressed in the females by modifier genes, when the trait becomes deleterious to females. Thus, sexually antagonistic selection on a trait and sex-specific gene expression can lead to the evolution of sexual dimorphism. Tusk is sexually monomorphic in the probocideans that are ancestral to both the African (Loxodonta africana) and Asian elephant (Elephas maximus). Tusk continues to remain monomorphic in the African elephant but has become sexually dimorphic in the Asian elephant. Tusk, therefore could be a sexually selected male trait that evolved according to the Fisherian model. Intriguingly, tuskless males occur at very high frequencies in some Asian elephant populations. The tusked and tuskless male morphs could be alternate male mating strategies, occurring at evolutionarily stable frequencies. Alternatively, the observed male tusk dimorphism, could be a consequence of artificial selection against tusked individuals, due to selective harvest of tusked males. Furthermore, male African elephants are more intensely poached for ivory than female elephants. Yet the frequency of tuskless individuals has increased more rapidly among females than in males. In essence, sexual dimorphism could be evolving among such poached populations. Is such rapid, contemporary evolution of sexual dimorphism, possible through the Fisherian modifier gene mechanism? A 2-loci genetic model (with X-linked trait gene and an autosomal modifier gene) (Rice 1984), a slight variant of the model (with X-linked modifier gene, and an autosomal trait gene) and an entirely autosomal model, were analyzed for the rate of evolution of sexual dimorphism, under different selection pressures for tusk possession. Negative frequency dependent selection was introduced into the model of tusk evolution in accordance with Gadgil’s model for the evolution of male dimorphism as consequence of sexual selection (Gadgil 1972). In two of the 2-loci models (in which tusk gene in autosomal), tusklessness evolved much more rapidly in females than in males, under equal negative selection pressures. The models predict several combinations of time-lines and negative selection pressures for effecting a particular change in the frequency of tusklessness. Model predictions were com-pared with observed changes in the frequency of female tusklessness, in one South Ugandan, African elephant population (∼2% to 10% in 5 to 9 generations) and male tusklessness (∼5% to 50% in 25 to 40 generations) in one north eastern Indian, Asian elephant population. The models predict strong selection pressures of 30% to 50% reduction in fitness, that can effect an 8% increase in tusklessness, in the African elephant population, within time-lines of 9 to 5 generations (∼225 to 125 years) respectively. For the male Asian elephants, natural selection against tusked males on an already sexually dimorphic population, must have been in operation and shifted the population to 5% male tusklessness. The models predict that artificial selection with 20 to 30% fitness cost to tusked males, operating for 40 to 25 generations (∼1000 to 600 years) respectively, can further shift the population from ∼5% to ∼50% tusklessness. Asian elephant populations may already have been in a transient phase of evolution, tending towards tusklessness, with recent artificial selection hastening the process. The two major pre-dictions from this modeling exercise are (1) artificial selection could have played a significant role in the evolution of male tusk dimorphism in the Asian elephant (2) a lack of or very mild current sexual selection on tusks in the male Asian elephant. Both these predictions may be empirically verified. Chapters 2 and 3 are attempts at empirical verification of prediction (1) and Chapters 4 and 5 of prediction (2). From historical references to elephant harvest in Assam, we do know that artificial selection has been in operation, but whether it has played a major role in causing male tusk dimorphism needs to be established. It may be possible to detect signatures of significant past harvest from current demographic structure of an elephant population. Sustained biased harvesting of a particular sex and or age class from an animal population alters the sex ratio and age structure (relative proportion of individuals in each age and sex class) of a population considerably (Sukumar 1989). It may be possible to back infer the harvest scenario by studying the deviation of current age and sex ratios from natural age and sex ratios. In Chapter 2, I explored models of population dynamics under different harvest regimes and its effect on age and sex ratios. I described a method to infer unknown harvest rates and numbers from age and sex ratios, namely, adult female to male ratio, male old-adult to young-adult ratio, and proportion of adult males in the population using Jensen’s(2000) 2-sex, density-dependent Leslie matrix model. The specific combination of male and female harvest rates and numbers can be deter-mined from the history of harvest and an estimate of population size. I validated this model with published data on age and sex ratios of one Asian and African elephant population with fairly reliable data on elephant harvest as well. In Chapter 3, I applied this model to the demographic data that I collected from a wild Asian elephant population in Kaziranga National Park, Assam, India (where more than 50% of the adult males are tuskless). Male polymorphism of sexually-selected male traits occur at stable frequencies in populations of several species. The different male morphs of the trait are hypothesized to be alternate male mating strategies with equal life time reproductive fitness. Male Asian elephants of Kaziranga National Park, Assam are dimorphic with respect to tusk possession: ∼50% of the males are tuskless (and are locally called makhnas). Makhnas could be trading tusk for either longevity, larger body size, testicular volume and or duration of musth as alternate mating strategies. On the other hand makhnas may have increased to a very high frequency primarily due to selective removal (captures for domestication and hunting for ivory) of tusked males from the population for centuries. The aim of Chapter 3 was to examine the role of artificial selection in the evolution of makhnas. Prolonged male-biased harvest(removal from the population) is bound to alter the demographic structure of the population and leave a signature of the intensity and type of harvest on the residual population structure. The Kaziranga elephant population was considered as representative of elephant populations of north east India; A harvest modeling approach (described and validated in Chapter 2) was used to infer unknown harvest of elephants from demographic parameters estimated by sampling this elephant population during 458 field days in the dry season months of 2008–2011. The Kaziranga elephant population appears to have been harvested approximately for the past 700 to 1000 years with adult tusked males being harvested at approximately twice the rate of adult tuskless males, adult females and their immature offspring of both the sexes. The currently observed high frequency of tuskless males in Kaziranga therefore, may be a consequence of sustained artificial selection against tusked males for several centuries. The previous two Chapters have only examined some mechanisms for the loss of tusks in elephants. I proceeded to examine the possibility of evolution of tusks through Darwin’s mechanisms of male-male competition for mates and female mate choice. Elephant tusks are cited as an example of a male trait that has evolved as a weapon in male-male combats. In Chapter 4 I examined the role of tusks in establishing dominance along with two other known male–male signals, namely, body size and musth (a temporary physiologically heightened sexual state) in an Asian elephant population in northeastern India with equal proportions of tusked and tuskless males. I observed 116 agonistic interactions with clear dominance outcomes between adult (>15 years) males during 458 field days in the dry season months of 2008–2011. A generalized linear mixed-effects model was used to predict the probability of winning as a function of body size, tusk possession and musth status relative to the opponent. A hierarchy of the three male–male signals emerged from this analysis, with musth overriding body size and body size overriding tusk possession. In this elephant population tusk possession thus played a relatively minor role in male–male competition. An important implication of musth and body size being stronger determinants of dominance than tusk possession is that it could facilitate rapid evolution of tuskless males in the population under artificial selection against tusked individuals, which are poached for ivory. If not a weapon, tusks could be a male ornament that female elephants find attractive. I explored the interplay of the three male traits (body size, musth and tusk), male mating strategies and female mate choice in Chapter 5. In some species males obtain mating opportunities by harassment of females. Given the striking size difference between an adult male and female elephant, with males weighing at least 30% more than females, male coercion of females to mate is a possibility. A detailed study of the courtship behavior revealed that overt male harassment of females is rare and the ability of a male to mount and stay mounted on a female for copulation is under female control. Therefore female Asian elephants can exercise choice to mate but this is subtly different from exercising mate choice itself. Age-related male mating strategy (reported for the first time in the Asian elephant) exists in the Kaziranga elephant population and this strategy limits the ability of females to exercise choice. Young males (<25 years) predominantly show a sneak mating strategy. Middle-aged males (25–40 years), when in musth, mate–guarded oestrous females from sneakers and attempted mating but sometimes resorted to sneak mating when out of musth. Old males (> 40 years) attempted mating only during their musth phase and were seldom sneakers. Large/musth males received positive responses from estrous females towards courtship attempts significantly more often than did small/non–musth males. Tusked non–musth males attempted courtship significantly more often than did their tuskless peers, and had a higher probability of receiving positive responses than did tuskless males. A positive response, however, may not translate into mating because of mate–guarding by the dominant male. Females permitted large/musth males to stay mounted significantly longer than small/non-musth males. Musth and large body size may be signals of male fertility. Female mate choice in elephants thus seems primarily for traits that signal direct benefits of assurance of conception. Tusked males may attain sexual maturity faster than tuskless males. Therefore it is worth exploring if tusks function as signals of male fertility when males are young (15 to 25 years); this may be possible through hormonal and behavioral profiling of young tusked and tuskless males from 10 to 20 years of age. Overall all musth and body size appear to play a larger role in enhancing male mating success than tusks. Tusked males appear to have a weak sexual selection advantage (male-male domi-nance and female preference) over their tuskless peers, only in the young age class (15 to 25 years) in this population. Males in this age age class, seldom come into musth that would over-ride tusk as a signal of male dominance. Current sexual selection on tusks in this population, appeared to be insignificant and this may be verified through genetic analysis of paternity success. An important implication of musth and body size being stronger determinants of mating success than tusk possession is that, it could facilitate rapid evolution of tuskless males in the population under artificial selection against tusked individuals, even in a slow breeder such as the elephant. Musth may have evolved much later than tusks in elephants, therefore it is possible that tusks evolved under sexual selection before musth evolved. However, body size, in mammals in gen-eral appear to be under both natural and sexual selection. Gould has shown that the absurdly large and palmate antlers of the extinct Irish elk, scales allometrically with body size (Gould & Lewontin 1979). Phylogenetic studies of elephant evolutionary radiation indicate a general trends towards increase in body-size with size reduction and tendency towards dwarfism occurring only in island habitats (Palombo 2001). Tusk development, which is essentially tooth development may be closely linked to cranium development. Cranium development in turn may be linked to body size through allometric scaling laws. If so, any selection on body size is bound to act on tusk size. I propose that the evolution of elaborate tusks seen in elephants is primarily due to natural and or sexual selection acting on body size, and tusk just hitched a ride with body size. Tusks may be maintained in spite of tuskless males occurring in the population only because of a rather weak sexual selection advantage to tusk possession in contests in which males are symmetrical with respect to body size and musth status.

This study looks at the impact of elephant feeding on the Xeric Succulent Thicket component of Eastern Cape Subtropical Thicket (ECST) in Addo Elephant National Park (AENP). Observations of elephant feeding were carried out and vegetation transects were surveyed for impact of elephant feeding. The results indicated that the Nyati elephants spent the majority of their time grazing (nearly 90%), particularly the cow-young herds, and especially when the herd gathered in larger numbers. Browsing events were concentrated on Acacia karroo (81%) and there was no significant difference between the sexes in their preference for this species. Despite being subjected to most of the browsing, the majority of A. karroo trees were undamaged and the effect of elephants was generally light. It appears unlikely that, three years after re-introduction to Nyati, the elephants have had an effect on community structure of the vegetation. Surveys were conducted on stands of the alien invasive weed prickly pear Opuntia ficus-indica, and it was recorded that elephants in Nyati have had a dramatic effect on prickly pear, utilising all adult plants assessed and destroying 70% of them. This level of destruction in such a short period of time suggests that prickly pear is a highly favoured species. The results from the present study suggest that elephants can play a role in the control of prickly pear. Results are discussed in terms of elephants as both megaherbivores and keystone species, and as agents of intermediate disturbance.

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.