A crucial question in the debate about reintroducing elephant culling is whether the long-term effects of elephants and mesobrowsers on woodlands are similar. Sufficiently high biomass-densities of mesobrowsers may, following reduction or removal of elephants, continue to heavily impact earlier life-history stages of a similar suite of woody plants that elephant impacted, preventing these species from maturing. Thus a similar end-point for woodland structure and composition is achieved. No study exists in the literature where woody plant and habitat utilisation of the savanna browser guild has been determined in the same locality over the same period. A review of 49 years of literature implied that the two groups impact the same core woody-species in the same habitats. Dietary and habitat utilisation of guild members was determined in Hluhluwe-iMfolozi Park, South Africa. A small suite (n = 8) of woody species formed the core diet of all guild members. Herbivores’ densities were determined using a novel GIS approach; all members of the guild showed extensive overlap in habitat use. GPS collars and a GIS were then used to detect zones of different density of impala in the landscape, thus defining, for the first time, a natural fine-grain browsing gradient. Densities of woody seedlings were significantly less (average 48% reduction) in areas of high versus low impala density. A simple browse-browser model, incorporating, in a novel approach, functional groups of plant species, was parameterised from these results and an extensive review of the literature. Outputs suggest that over the long-term (100 years), impala will have a similar impact on woodland structure as elephant. An apparently strong synergistic effect between impala and elephant impact, suggests that reduction or removal of either impala or elephant will radically reduce long-term destruction of woodlands. In smaller or medium sized reserves, where control of mesobrowser populations is practical, profitable and more acceptable than elephant culling, these findings imply a re-direction of management efforts. Management should consider the biomass-density of both groups, rather than just focus on the system’s perceived ‘keystone’ species. Such principles may also apply to temperate and other systems.

A Research Report submitted to the Faculty of Science, University of the Witwatersrand, Johannesburg, in partial fulfilment of the requirements for the degree of Master of Science. Johannesburg, 25 May 2015. / The African elephant, Loxodonta africana displays sexual segregation, a phenomenon which describes males and females of the same species living separately, except during the mating season. Despite it occurring in many sexually dimorphic species, the factors that govern sexual segregation are still poorly understood. The aim of my study was to investigate whether or not African elephants in the 1825 km2 Associated Private Nature Reserves (APNR), Limpopo Province were sexually segregated as a result of habitat segregation. I tested the Forage Selection Hypothesis (FSH) which, based on the Jarman-Bell principle, predicts that smaller females are more selective foragers as a result of their high energy demands and poor digestive capabilities in comparison to the larger males. Using the GPS location data of 18 collared adult elephants (12 male and 6 female) from November 2008 to November 2010, I plotted both the total (95% isopleth) and core (50% isopleth) home ranges of individual elephants. I used these home ranges to i) confirm sexual segregation in the APNR, ii) determine whether or not there was a difference in vegetation composition of the home ranges between males and females, and iii) to establish how frequently male and female elephants were associated with each of the vegetation types located within their home ranges. All analyses were done at both the total and core home range level. Home range overlaps were rare between male and female home ranges, particularly at the core home range level, confirming sexual segregation of elephants in the APNR. The vegetation composition data of the home ranges as well as the frequency of association by elephants with each of the available vegetation types (using GPS locations) revealed no significant difference between male and female elephants. Therefore, habitat segregation did not explain sexual segregation by elephants in the APNR. I propose that future studies should consider: i) temporal distribution of elephants to assess how much time each sex spends in each of the available vegetation types; ii) other resources, particularly the availability of water, in addition to the availability of forage, since water limits elephant movements; and iii) social segregation in conjunction with habitat segregation, since elephants display sex-specific differences in social organisation. A comprehensive understanding of the factors that govern sexual segregation of elephants might contribute to conservation management of elephants in the APNR and other small reserves.

A thesis submitted to the Faculty of Science, University of the Witwatersrand, Johannesburg, in fulfilment of the requirements for the degree of Doctor of Philosophy. 5 September 2016, Johannesburg, South Africa. / There is an urgent need to study the effects of elephants on biodiversity given the ability of megaherbivores to transform vegetation composition, structure and function by killing selected plants. Within a biodiversity framework of different aspects of diversity across different scales, we need to understand elephant effects across time and space, acknowledging disequilibrium dynamics of savannas. However, most savanna studies are conducted either over a short time frame, over a limited spatial extent, or without species compositional data. The Linyanti riparian woodland in northern Botswana represents a valuable opportunity to study the effects of elephants as it is subject to extremely high elephant concentrations in the dry season as elephants congregate on the perennial river. Moreover, because of trampling effects by large herbivores and high soil moisture, fire is largely excluded, allowing the study of intense elephant impacts in relative isolation. This PhD thesis aims to assess long-term (16-18 years) compositional and structural change at a large spatial scale (50 km of riverfront) of the Linyanti riparian woodland, built upon two earlier studies in 1992/2 and 2001. Specifically, it aims to establish the effects of elephants on 1) the spatial heterogeneity of disturbance across the woodland; 2) compositional changes of the canopy tree layer caused by elephant impacts; 3) the potential of the woodland to regenerate from seedlings; 4) structural changes due to woodland decline and shrub increase. It finally aims to synthesise these findings for biodiversity and the implications for conservation and management. Spatial heterogeneity was assessed by delineating patches of intense disturbance using the clustering algorithm DBSCAN. I manually marked dead trees within a 2000 ha overlapping riparian area from the 1992, 2001, and 2010 aerial photographs and determined these trees were significantly clustered in the landscape to form patches of disturbance. Disturbance patches were highly dynamic over the period where small patches appeared, grew and coalesced over time, whilst a few patches fragmented or disappeared. The overall dynamic was of smaller patches coalescing resulting in the total patch area increasing from 6% in 1992 to 23% in 2010. Mortality increased mostly in the inter-patch areas but the overall dead tree appearance rate of 0.28 trees.ha.yr-1 was not much higher than a background tree death rate calculated for exclosures in other areas. The slow mortality rate coupled with progressive decline suggests there was little recruitment into the canopy to replace the trees that were lost. Even though large areas remained that were not classified as disturbance patches, there was evidence of increased fragmentation where inter-patch areas became increasingly small and isolated. This increase in greater areas of disturbance represents a state shift to decreased heterogeneity although landscape patchiness still remained in 2010. Projections were that mortality rate and patch formation would decrease. To assess compositional changes, I reconstructed the pre-1992 canopy tree woodland by combining both living and dead trees in 1992, and compared this to the 1992 and 2008 woodland composition. The woodland showed progressive declines from an Acacia spp.-Colophospermum mopane dominated tall tree woodland pre-1992 to a woodland in 2008 composed primarily of two resilient species (C. mopane, Combretum hereroense), and one avoided species (Philenoptera violacea). I compiled Size Class Distributions of individual canopy tree species and compared proportional high impact on living and dead trees between 1992 and 2008. High elephant impact was defined as more than 50% stem circumference ringbarked or with the main stem or majority of side stems broken. I found that elephant impact was the likely cause of the woodland decline, although wind and natural senescence were variably important for some species. The acacias had nearly disappeared from the woodland, declining in proportional abundance from 30% in the reconstructed pre-1992 woodland to just 4% in 2008. Over time there was a progressive shift in elephant impact from abundant preferred and vulnerable species like Acacia spp. and Terminalia spp. to species more resistant to debarking like Combretum imberbe and Berchemia discolor. The abundant species C. mopane proved highly resilient to intensive elephant impact. The seedling layer (plants below 0.5m) had high proportions of canopy tree species including the acacias, and all but the rarest species were recorded. This suggests regeneration of the woodland is possible but there was a demographic bottleneck of seedling mortality with few saplings recorded over the time period. To determine the structural changes which have taken place, I separated shrub species and canopy-forming tree species and assessed density changes in the sapling (<2.5m) and tree (>2.5m) layers. Tall (>2.5m) canopy tree density decreased by half between 1992 and 2008, representing an annual loss rate of 2.7% without replacement. Except for Colophospermum mopane, there was no compensatory regeneration in the form of saplings. Colophospermum mopane was highly resilient to elephant impacts, coppicing vigorously following impact to form local ‘browsing lawns’ which may benefit other browsers. The overall shrub density increased 2.5 times while one shrub species (Combretum mossambicense) increased five-fold in density and came to constitute 50% of the total woody plant density. This shrub species increased rapidly, at an exponential growth rate of 10.5% per year, representing pervasive shrub encroachment. Its invasion wave was incipient in 1992 and by 2008 many of these plants had grown beyond 2.5 m in height, forming a dense screen. Small plants of this species <1 m in height had become sparse by 2008, suggesting that the invasion had become curtailed by then. I proposed that the spread of this shrub was due to its unpalatability by elephants, although it is an important browse species for ruminants. A potential global driver of enriched atmospheric CO2 or regional aridification could not be ruled out. The state shift from woodland towards dense shrubland caused by differential elephant impacts has potential negative consequences for structural and functional diversity. I attempted to synthesize the findings for biodiversity and concluded that there was a state shift towards pervasive disturbance with a corresponding decline in spatial heterogeneity, although composition of the disturbance patches was not studied. There has however, not been a state transformation from woodland and stands of tall trees were still present in the woodland. Coupled with the potential regeneration of the woodland from seedlings, these findings highlight the importance of long-term studies of non-equilibrium savannas. The main threat to biodiversity of the woodland was not elephant-induced mortality of large trees, but rather the lack of recruitment and the pervasive shrub encroachment of a single species. It may be, however, that alternate states of canopy trees and unpalatable shrubs exists, enhancing long-term functional diversity, provided the system remains relatively open and elephants are free to move to other areas. Ultimately the only management strategy of relatively open areas with high elephant concentrations is to accept changes and support transfrontier conservation efforts. I further assess the limitations of this study, and make recommendations for future study, specifically highlighting the need for a longer-term palaeo-ecological study to evaluate compositional changes due to episodic recruitment events. / LG2017

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.

Perceived predation risk alters animals’ behavior. This shift in behavior often comes at the cost of attaining resources. Generally, African elephants (Loxodonta africana) experience little predation pressure; however, the risk of predation by lions (Panthera leo) increases other prey species are less abundant. In elephant herds, related females and their offspring travel together in family groups, led by the eldest female. Response to predation pressure was examined by playing lion calls to the population of 437 elephants at the Main Camp Section of Addo Elephant National Park (AENP) in South Africa. Unfamiliar lion calls from a single male and two males, static, and running water were played from a remote speaker to elephants at waterholes. These trials were recorded by video. Behaviors of elephants were then extracted from video into focal observations of thirty second segments before, during and after a sound was played. I analyzed these data using parametric t-tests and non-parametric randomization tests. When no sound was played, elephants did not alter their behavior. Water elicited low levels of distress behaviors. Elephants behaved in a threatened or annoyed manner toward static. Elephants changed their behavior more in response to lion calls than to the controls, namely by decreasing drinking and increasing walking and distress behaviors. I also examined how individuals differed in their responses to the lion calls based on a number of demographic factors. Adult and subadult females performed more social behaviors after lion calls when the matriarch was absent than when she was present. Furthermore, when group size was larger and more calves were present, females decreased drinking and increased time exhibiting distress behaviors. Based on this and other studies it can be concluded that elephants of different demographics perceived similar levels of elevated risk when hearing lion calls. Landscape of fear models are useful for assessing habitat use by prey species in response to real and perceived predation risk. The present study corroborates findings from a study in East Africa that elephants perceive threat from lions based on calls alone and appear to distinguish levels of threat by the number of lions calling.

In this thesis, I explore the June 1946 issue of Dance Index: Joseph Cornell’s “Clowns, Elephants and Ballerinas.” Through the archive of materials collected and presented by Cornell, I attempt to understand the histories of performance offered to the magazine’s readers. Despite the rich field of scholarship dedicated to Cornell and his art, very little work has been dedicated to his contributions to Dance Index. I interpret “Clowns, Elephants and Ballerinas” as both a collage and a series of histories, and I present the magazine as a serious work in Cornell’s oeuvre. I also endeavor to provide an understanding of Cornell’s working method, his sense of history, and the ways his juxtapositions of word and image provide meaning to readers. Weaving together the visual and textual, contemporary and historical, Cornell explores performance legacies, American and European exchange, and pantomime, dance, and circus performance tradition through this magazine issue. Cornell uses each of his diverse materials to explore larger social and political issues as well as artistic traditions. “Clowns, Elephants and Ballerinas” represents a crystallization of a moment in one of his many “explorations.” / text

This thesis is an investigation into, and a gathering of evidence on, the various ways in which two iconic species, whales and elephants, and the two conventions which govern their management, the 1946 International Convention for the Regulation of Whaling (ICRW) and the 1973 Convention on International Trade in Endangered Species (CITES), are linked in international law and politics. After explaining the nature of international conventions governing wildlife species generally, the respective histories of the two conventions are considered: first, that of the ICRW is considered, together with its strengths, weaknesses and current position; after which a similar assessment is made of CITES. The history of linkage between the two is considered, including attempts made to use the one to undercut the other. Various aspects of the protection, use and management of the two species are then canvassed; and it is shown how important political actors hold apparently mutually exclusive views. Throughout, the position of South Africa is particularly considered. The importance of protecting biological diversity is then considered, together with the potential harmonising role of the 1989 Convention on Biological Diversity (CBD), and the political stances of various countries, together with ongoing analysis of efforts to effect change. The natures of whales and elephants as symbols, and as special animals, are then considered. In conclusion, it is explained that both treaties could work if the political drive was present - but that this is currently absent, and the environment is suffering whilst politicians argue over the best courses to follow to protect natural resources. It is suggested that the reason that the arguments in respect of whales and elephants, the ICRW and CITES, are so bitter is because so much is at stake - for the fight on this battleground is not simply about the particular species, but the course the world as a whole should follow in all of its use of natural resources. Understanding the links between species and between treaties helps us to understand alternative possible courses. By exploring one such set of links that has not previously been analysed, the research presented in this thesis is intended to make a contribution to that understanding (both internationally and within South Africa). / Thesis (LL.D.)-University of KwaZulu-Natal, Pietermaritzburg, 2008.

The continual increase in elephant numbers across southern Africa raises concern, though the interventions to manage these populations are more contentious. Within small, enclosed reserves active management is necessary to limit elephant impact. Various management tools exist with which to control fluctuations in elephant population numbers or density and simulate natural large stochastic events to control population growth. During this study, undertaken in the Munyawana Conservancy, KwaZulu-Natal, and Pilanesberg National Park, North West Province, South Africa, several management options were implemented. In order to lower the population numbers, family groups were translocated from the Munyawana Conservancy to other reserves, while to reduce population growth rate an immunocontraception was implemented. Both conservation areas introduced older bulls to normalise the bull population age structure, and expanded the conservation area by inclusion of new land to reduce population density. The influence of these management interventions on the elephant population were measured by their social, behavioural, spatial and movement responses. The older bull introduction was successful as bulls set up exclusive bull areas. There was a quick, subtle affect on the bull groups' size immediately after the older bull introduction, while there was no immediate change within the resident bulls' musth behaviour or duration. During area expansion, elephants appeared to perceive the new unexplored area as a threat although this threat became reduced through time as they became more familiar with it. The spatial scale of response was relatively small, while the temporal scale of response was relatively large. Rotational immunocontraception was shown to be a successful tool to alter herd structure by aging the population and maintaining a low population growth rate. The process of immunocontraception darting had no significant effect on herd associations and movement rates, accordingly the duration of the disruption effects were short lived. During multi-management interventions, no differences were found within the elephant social grouping. Management interventions may pose unforeseen social risks and different populations may respond differently to management induced stress. Therefore, interventions need to be considered for each elephant population which will achieve the conservation area's objectives with the most effective outcome, but with lowest holistic impact. / Thesis (M.Sc.)-University of KwaZulu-Natal, Westville, 2012.

Increasing human population and development in Africa restricts land and resources for African elephants (Loxodonta africana) and threatens the preservation of the species. Despite the importance of conservation for many governments in Africa, many local communities have negative views of elephants and exhibit anti-conservation behavior. By looking at the history of wildlife policy in Tanzania, this paper seeks to understand these opinions through a historical context. Three case studies of conservation initiatives were evaluated to determine what aspects of conservation initiatives promote long-term pro-conservation behavior on the part of the local community. Ultimately, conservation initiatives should establish a framework whereby local communities are empowered through conservation.