The elephant debate deals largely with population size, how elephant numbers change over time, how they may affect vegetation, and how their populations should be managed. Trends in elephant numbers frequently motivate management decisions, and past efforts to alleviate elephant impact aimed at controlling population size. However, methodological and statistical constraints may influence interpretation of trends and lead to incorrect management decisions. Furthermore, inferences about the response of elephant populations to specific management actions are seldom based on scientific evidence. In this thesis I assess the consequences of survey design and monitoring features on the interpretation and statistical reliability of population trends as well as the effect of population management on elephant densities and population growth rates. To do this, I collated information on elephant population estimates and past management actions across Africa. I used information from the northern Botswana elephant population to clarify temporal trends in elephant densities and numbers. Elephant numbers in northern Botswana increased from 1973 to 1993 while densities remained relatively stable. This difference in trends is due to an associated increase in survey area during the same time. In contrast, from 1996 to 2004 surveyed areas remained constant in size and neither elephant numbers, nor densities changed significantly during this time. This apparent stabilisation in numbers may have resulted from density-related elephant dispersal. This case study suggests that in open populations movements may complicate the interpretation of trends, and that differences in the rates of change in numbers and densities may have different management implications. The precision of population estimates, sample size, population size, and the magnitude of the annual rate of population change to be detected, affect power to identify trends. Two-thirds of the 156 time series that I assembled apparently were stable, and only 30 % of these had sufficient statistical power to detect population changes. These apparent stable trends without sufficient statistical power are inconclusive and should not be used to inform management decisions. Past elephant population management practices may have increased densities and growth rates in African elephant populations. Case studies of populations that were exposed to different management actions indicated that fencing of populations and water supplementation may have enhanced growth rates probably by influencing dispersal patterns. Thus, past management practices may have contributed to the ‘elephant problem’ by enhancing local elephant densities and population growth rates. In this thesis, I showed that trends based on elephant numbers may be misleading when the area over which elephants were counted, increased in size. Second, despite much effort and resources devoted to the monitoring of elephant populations for more than 50 years, population estimates and time series including such estimates had low quality, thereby reducing statistical power to detect trends in population change. Third, population growth rates were associated with management, where elephant population densities grew at faster rates when managed. Future conservation efforts should take into account the methodological and statistical constraints that may influence trend analyses of elephant populations and take cognizance of the fact that management decisions need to be evaluated against expected outcomes. Copyright / Dissertation (MSc)--University of Pretoria, 2009. / Zoology and Entomology / unrestricted

Thesis (M.S.)--University of Pretoria, 2007. / Title from PDF title page (viewed on Nov. 12, 2008). Includes bibliographical references.

The successful conservation management of African elephants depends largely on understanding the fundamental processes driving the population regulation of this species. Southern Africa’s increasing populations have raised concern over the impact of high elephant densities on the system, in stark contrast against the elephant’s more precarious position in other parts of Africa. As we search for solutions from the processes of historical elephant regulation, we realise that there is a decided lack of empirical evidence to explicitly direct our efforts. In this PhD, I attempt to investigate the application of the classic pattern of large herbivore population limitation, which mainly involves high juvenile mortality in response to stochastic environmental events, to African elephant population dynamics. Firstly, I evaluated the magnitude and frequency of mortality events that would be required to prevent elephant population growth. The death of 85 % of infants and weaned calves would need to occur twice a generation, while a single severe mortality event (causing the death of all infants and weaned calves and 10 % of the rest of the population) once a generation would be sufficient. However, the severity of these events is not matched in natural occurrence in Africa today and only a single recorded event in Tsavo National Park, Kenya, in the 1970’s has come close when more than 7 000 died during a very severe drought. Secondly, I evaluated the potential role of fire as a stochastic, massmortality event limiting elephant populations. I found that fire functions in a similar manner to other environmental catastrophes and primarily causes high juvenile mortality. However, this catastrophic event also highlighted the extreme behavioural and physiological impacts experienced by the elephant population involved. The potential role of these types of events on long-term female fecundity needs further investigation. In isolation, this type of mortality event would need to occur with high frequency to prevent population growth. However, in combination with a decrease in female fecundity, these stochastic events may have a much greater impact on population demography than first thought. Thirdly, I investigated a potential mechanistic link between stochastic mortality events and juvenile susceptibility to resource limitation. Allometric relationships dictate that juveniles select a diet of higher quality than adult elephants. We found that this was achieved by weaned calf selection of higher quality plant parts, although use of plant types and plant species was similar to that of adult females, who they move across the landscape with. The strong sexual dimorphism exhibited by this species was reflected in adult male use of lower quality forage than adult females (or juveniles) in both dry and wet seasons. Diet quality scaled negatively with body size, but adult females consistently selected a higher quality diet than adult males, irrespective of body size. The nutritional and reproductive demands placed on an individual during different life-history stages therefore influence foraging strategies, together with nutrient requirements, e.g. phosphorus for pregnancy/lactation selected consistently by females when unrestricted in the wet season, protein for growth selected consistently by weaned calves. Competitive displacement of adult females to feed at higher levels in the canopy by calves also influenced feeding behaviour. Therefore intraspecific body size, nutritional requirements (in terms of nutrients and energy) and competition had a strong influence on foraging strategy employed by age-sex classes of elephants in response to seasonal environmental change. More selective juvenile foraging requirements means that juveniles are most susceptible to resource limitation, for example during stochastic environmental events such as droughts. In small, closed systems, juvenile mortality is likely to have a strong influence on elephant population regulation, with a slight, temporary decrease in female fecundity possibly acting in conjunction with juvenile mortality effects. Therefore, stochastic environmental events such as drought and fire may be the only natural incidence of population regulation to occur in these systems, where populations continue to grow exponentially and there is no evidence of density-dependence (as in the case of many small, fenced reserves in South Africa). In large, open, high-density systems in other parts of southern Africa, density dependence acts strongly on female fecundity and causes low levels of juvenile mortality in areas of local population aggregation. Therefore, in isolation, natural juvenile mortality is unlikely to regulate African elephant populations, but in conjunction with decreased female fecundity in response to density-dependent feedbacks and stochastic environmental events, population regulation may occur. The management of long-lived megaherbivore species with similar demographic drivers must include an appreciation of the complexity of population response to manipulation of mortality or fecundity effects. Small changes can potentially result in large shifts in population dynamics. Further insight into the mechanisms driving these processes will allow sound scientific support of megaherbivore management decisions to be made throughout Africa. / Thesis (Ph.D.)-University of KwaZulu-Natal, Westville, 2008.

This study presents an investigation into the population dynamics and social structure of a small, closed elephant population. Specifically, it examined population growth rates for evidence of density-dependent regulation. It also quantified the association patterns of female elephants groups, and male elephants groups. Social structure was examined using Hamilton’s kinship theories of inclusive fitness, and age. Male-female patterns of association were also examined for inbreeding avoidance behaviours. The study population was located in Addo Elephant National Park, South Africa. Density-dependence was assessed using a long-term data set. Densities were considerably higher than estimated carrying capacities. Population growth rate was positively correlated with increasing density. No relationship between birth rate, the age of first calving or calf sex ratio and elephant density was detected but there was a positive relationship between birth rate and rainfall during conception year. Mortality rates, particularly for juveniles, were low, and mean inter-calf interval was 3.3 years. There is no evidence of density dependent regulation in this population. These findings indicate that density dependence should not be considered as an option in the control of elephant numbers in this Park, or where elephant resources are not seasonally limited. Examination of association patterns of the adult female component revealed that associations were not random at the population, family or individual scale. This is the second study on African elephants to confirm previous behavioural studies that predicted that preferred associates were close maternal relatives. This supports many studies showing that social species preferentially associate with their kin. The adult males in this population were found to have a well differentiated society with non-random associations. Generally, males were found to have weak associations with most other males and strong associations with only a few males. This association pattern was found to be persistent over the time frame of the study, as indicated by the time lag analysis. Males returned to their natal family, even when maternally related females were in oestrus. Oestrous females directed positive behaviours towards musth males. It appears that behavioural inbreeding avoidance mechanisms in this small, closed population are inhibited: musth status seems to override inbreeding avoidance. General principles from this case study were interpreted in terms of their applicability to other small, closed populations.