abstract: Prior to the COVID-19 global pandemic, ecotourism represented the tourism industry’s fastest growing segment with projections estimating that ecotourism would become the world’s largest tourism type by 2030. While the tourism industry will need several years to rebound, if historic trends tell us anything, it is that ecotourism will continue to represent a large portion of the overall industry and will continue to grow at a rate that outpaces all other tourism types. In theory, ecotourism promotes sustainable socioeconomic development while also minimizing negative environmental impacts. Unfortunately, research suggests that this is not always true, and many examples exist of ecotourism causing more harm than good. In order to combat these potential negative impacts, the ecotourism industry has become increasingly reliant on ecotourism certification programs to act as an assessment tool that identifies ecotourism’s best practitioners while minimizing false advertising present within the industry. Despite these beliefs in the efficacy of certification, there is a lack of empirical research to actually support certification as an effective assessment tool. Furthermore, little research has been conducted that assesses the impacts that certification itself has on ecotourism businesses (both certified and uncertified) and the local communities dependent on ecotourism. My dissertation employs a mixed methods design and combines qualitative and quantitative research methods spanning multiple geographic scales to develop an understanding of certification programs as they exist today and to discern the impacts that certification itself may cause for all those either directly or indirectly involved in ecotourism. My findings ultimately suggest that certification reform is needed if certification programs are expected to be the assessment tool ecotourism experts claim them to be. Specifically, as certification exists presently, there is: no universal guideline or standard for existing certification programs to follow, a disconnect between the advertised benefits certification offers and the actual benefits received, and a lack of market penetration both amongst ecotourists and ecotourism businesses. Each of these must be addressed before certification can live up to its full potential. Furthermore, I found that certification may impact community socioeconomic dynamics, particularly by creating or exacerbating community wealth distribution. / Dissertation/Thesis / Doctoral Dissertation Biology 2020

A general and increasing biodiversity loss has been observed since the 20th century. Faced with the extreme rapidity of population declines, conservation biologists seek to understand the limiting and regulating factors driving changes in animal populations. This is particularly important for rare species as small population size increases extinction risk. Birds are amongst the most studied animals in this context. As a group that occupies a high trophic level, raptors are particularly vulnerable to external changes and are generally regarded as useful indicators of ecological change. The Black Harrier Circus maurus is an avian predator endemic to southern Africa, which breeds essentially along the South African coast within the Fynbos biome, and inland within the Karoo biome. Its population size has been estimated at less than 1,000 breeding birds, and the species is currently considered as Endangered in South Africa, Namibia and Lesotho. Although some studies have been conducted on Black Harriers in the last four decades, the reasons for its scarcity currently remain little known and insufficiently explored. Filling this knowledge gap is therefore essential for its conservation. In this context, the main goal of this thesis is to develop an overall comprehension of how various environmental factors may affect the breeding and health of this Endangered species, at both population and individual levels. I conducted my fieldwork during the 2012-2015 breeding seasons in two contrasting geographical regions: one along the west coast in the Western Cape Province, and the second one inland in the surroundings of Nieuwoudtville in the Northern Cape Province. For some chapters (Chapters 1-3), I analysed historical data collected by Dr. R. E. Simmons during 2000-2011 breeding seasons.

The Namaqualand area in the North Western Cape, South Africa is unique in comparison to other similar semi-arid areas of the world. It has a high biodiversity and endemism and is consequently an area of interest for a growing number of conservation initiatives. Climate plays an important role in influencing the phenology and growth of the vegetation in the area. Remote sensing techniques were used to reveal the vegetation patterns in the greater Namaqualand area and to relate them to climatic variables. To do this we used the normalised difference vegetation index (NDVI) to relate biomass to altitude, rainfall and vegetation type. Each vegetation type in the area had a unique temporal signature and the climatic variables influencing the summer rainfall and winter rainfall vegetation types differed significantly from each other. Mean annual NDVI was significantly correlated to precipitation and potential evapotranspiration (PET) (r = 0.60, -0.63 respectively). A multiple regression model explained 52% of the variance when Mean Annual NDVI was related to climatic variables. Mean NDVI in August (the month of maximum NDVI in most of Namaqualand) was significantly related to PET and the current plus two previous months of precipitation (r = -0.72 and 0.74 respectively). A multiple regression model for mean NDVI in August and climatic variables explained almost 58% of the variance. The results suggest that NDVI can be used successfully as a measure of growth and phenology in the Namaqualand area and that NDVI could be used in climate models, drought prediction, desertification predictions and a number of other applications in the future.

Their high energetic demands make seabirds sensitive to changes in prey availability, which is often reflected in their diet and energetic expenditure during breeding. Populations of the three seabirds endemic to southern Africa's Benguela upwelling ecosystem that rely on small pelagic fish have decreased dramatically over the last decade. In contrast, the population of the greater crested tern Thalasseus bergii has increased. To understand these conflicting trends, I investigated the foraging ecology and energy requirements of greater crested tern breeding in the Western Cape, South Africa. Diet was assessed by a novel non-invasive methodology developed in this study, using digital photography. More than 24,000 prey items from at least 51 different prey taxa were identified, with 34 new prey species recorded, revealing a high degree of foraging plasticity for this seabird. Greater crested terns rely mainly on anchovy Engraulis encrasicolus (65%), which averaged 84 mm long. Prey composition differed significantly between breeding stages, with anchovy especially dominant at the onset of the breeding period and the diet becoming more variable as the season progressed. Time-energy models for breeding terns were built based on activity budgets collected from non-invasive video-recordings and focal observations. Foraging trips were significantly longer during incubation than the chick provisioning stages, and feeding rates doubled from early to late chick provisioning. This study illustrated a steady increase in energy needs of adults throughout the breeding season, due to their increased foraging effort to meet chick energy needs. In comparison to other Benguela endemic seabirds that also rely on small pelagic fish, terns displayed substantially lower energy requirements at both individual and population levels. I also explored the benefits underlying interactions within mixed-species aggregations by investigating the costs induced by kleptoparasitism between mixed colonies of greater crested terns and Hartlaub's gulls Chroicocephalus hartlaubii and colonies with greater crested terns alone. Video-recordings coupled with focal observations showed that terns suffer direct costs to chick provisioning rates and indirect costs through energy expenditure in a mixed-species colony, suggesting that these breeding assemblages may be a form of parasitism rather than a mutualistic association. Despite the detrimental effects of interspecific kleptoparasitism, the marked foraging plasticity and low energetic requirements of greater crested terns, described in this study, coupled with specific life history traits such as low fidelity to breeding sites and extended post-fledging care, are key factors that allow this species to cope with changes in the availability and abundance of their main prey. Understanding species-specific behavioural responses to ecosystem variations in the Benguela upwelling system is vital for assessing the impact of commercial fisheries on seabird populations and fish stocks. Finally, the implementation of the method developed in this study, in long-term monitoring programmes, may provide crucial knowledge for conservation plans and key input to realising an ecosystem approach to fisheries management.

Seabirds are considered one of the most threatened groups of birds in the world. They face additional mortality both on their breeding islands from introduced predators and at sea by fishing fleets, as fisheries bycatch, as well as other human impacts. Seabird bycatch has negatively affected many seabird populations worldwide, with trawl, gillnet and longline fisheries considered the most destructive to seabird populations. Seabird bycatch from trawl and gillnet fisheries has been significantly reduced in recent years, but large numbers of seabirds are still killed annually by longline fisheries. Of the two types of longline fisheries (demersal and pelagic), pelagic longlining is considered the most harmful to seabirds as lines remain closer to the surface for longer periods than demersal longlining, and it is harder to weight lines to ensure rapid sinking beyond the depth they are accessible to birds. Seabirds are killed when they swallow baited hooks and consequently drown. Despite the large number of studies investigating seabird bycatch from pelagic longline fisheries, there remain gaps in our understanding of seabird bycatch from pelagic longline fisheries. This thesis addresses some of these knowledge gaps and makes recommendations as to how seabird bycatch from pelagic longliners can be reduced at both a local and global scale. Chapters 2 and 3 investigate the factors affecting seabird bycatch from pelagic longliners off South Africa, provide a summary of seabird bycatch from the fishery for the period 2006–2013 and quantify the structure of seabird assemblages associated with pelagic longline vessels off South Africa. This was achieved by analysing seabird bycatch data collected by fisheries observers as well as data from sea trials onboard pelagic longliners. Seabird bycatch by pelagic longliners off South Africa over the 8-year study period has been significantly reduced from the 8-year period (1998–2005), mainly driven by a significant reduction in seabird bycatch rates from foreign-flagged vessels, which are responsible for c. 80% of fishing effort off South Africa. Seabird bycatch rates from South African vessels still remain high, four times higher than the interim national target of < 0.05 birds per 1000 hooks. The species composition of seabird bycatch off South Africa is best explained by an understanding of the structure of the seabird assemblage associated with longline vessels. For most species, bycatch and attendance ratios were similar, but for some species such as shy-type and black-browed albatrosses there were large mismatches, likely caused by differences in foraging behaviour and foraging dominance hierarchies. In Chapters 4 and 5 the foraging ecology of the most commonly recorded bycatch species off South Africa, the white-chinned petrel (Procellaria aequinoctialis), is investigated. An understanding of the foraging ecology of commonly recorded bycatch species enhances our understanding of seabird bycatch and helps to improve the design of current and future mitigation measures. The year-round movements of white-chinned petrels from Marion Island were investigated with Global Location Sensors (GLS loggers) and GPS loggers. Adult white-chinned petrels undertake only limited east-west movements of, with all birds remaining between southern Africa and Antarctica. These results strengthen the theory that there is limited spatial overlap year-round between white-chinned petrel populations from South Georgia, the southern Indian Ocean islands and New Zealand sub-Antarctic islands, suggesting that these populations can be managed as separate stocks. The diving behaviour of white-chinned and grey petrels (P. cinerea), another common bycatch species in Southern Ocean longline fleets, were examined with the use of temperature-depth recorders (TDRs), deployed on birds from Marion Island and Gough Island. My study was the first to analyse TDR dive data from any Procellaria petrel, and recorded them reaching maximum dive depths of 16 and 22 m, respectively. Current best practise suggests that baited hooks be protected to a depth of 5 m by bird-scaring lines, but my results suggest this depth should be increased to at least 10 m. Although line weighting is a proven mitigation measure to reduce seabird bycatch from pelagic longliners, fishers have concerns that it will compromise fish catches, crew safety and operational efficiency. In Chapter 6 I analyse line weighting data from trials onboard three pelagic longline vessels, to address the concerns of fishermen. My results show that sliding leads can be incorporated into pelagic longline fisheries without compromising fish catch, crew safety or operational efficiency. I thus recommend that sliding leads be used on pelagic longline vessels fishing off South Africa. By incorporating studies investigating the factors affecting seabird bycatch, seabird foraging ecology and the efficacy of seabird bycatch mitigation measures, my thesis has broadened our understating of seabird bycatch from pelagic longliners and makes meaningful recommendations to further reduce bycatch, both locally and globally. Although seabird bycatch rates have declined off South Africa, through the use of a number different mitigation measures, they still remain higher than the South African national target and thus more work is needed to achieve this target. To reduce seabird bycatch from pelagic longliners to acceptable levels, studies from the world's various longline fleets needs to be considered and improved upon, with seabird conservationists and fishermen working together to achieve this goal.

Arid and semi-arid systems comprise one third of the earth's surfaces. The succulent karoo alone comprises approximately 1660 endemic species and is one of the world's hotspots in terms of diversity. Although these areas are of high ecological value less than 10% of arid systems are conserved. This study was conducted in Paulshoek a remote village in the Leliefontein area of the Namaqualand. Historically Paulshoek has been under continuous selective grazing since the 18th century. A fence line contrast is investigated in an area that was fenced three years ago and in one that has been exposed to heavy, continuous, selective grazing over the past century (both areas were fenced prior to fencing). The objective of this study was to investigate the effect of short-term rest on key vegetation demographic parameters as well as the physical characteristics of communal river systems. No significant differences in plant species richness were observed across the fence line. However, the numbers of plants of each species within the plots were significantly different, depending on presence or absence of grazing. The grazed sites had more species tolerant of grazing such as geophytes, annuals and toxic species such as Galenia africana. Heavy grazing in this ecosystem has resulted in increased proportions of unpalatable species with a slow reduction in the number of palatable species. Species cover was significantly reduced on the grazed sites; however, these reductions were insignificant for Lyceum cinereum species due to large standard deviations. Heavy grazing of palatable sedge species (Scirpus nodosus) resulted in significant reductions in the number of flowers of plants, in the grazed sites. Grazing not only reduced the size of the plants, it also reduced flowering, fruit set and recruitment rate. There was no significant difference in the physical and geomorphologic characteristics of the river across the fence-line. This study indicates that short-term rest has major implications for plant recovery. Educating communal farmers to use a farming system that insures temporary rest on certain areas of the rangelands at certain times of the year will go a long way in improving species richness in Namaqualand.

Populations of secondary tree-cavity nesting bird species are often limited by a shortage of natural nesting sites. For the Southern Ground Hornbill Bucorvus leadbeateri that typically nests in natural tree cavities, the shortage of nesting sites is one factor potentially limiting population growth. The species is listed as endangered in South Africa, and vulnerable throughout the rest of its range. Nest boxes can improve the conservation status of threatened birds that are limited by nest-site availability. However, nest boxes or other types of artificial nests are not always beneficial to the target species, and their value as a conservation tool needs to be tested for each species. Wooden nest boxes were installed for ground hornbills in a study area in north eastern South Africa with a paucity of natural nest sites. In this thesis, I assess productivity, timing of breeding, and dispersal in the Southern Ground Hornbill in a study area supplemented with nest boxes and discuss the implications for the conservation of this endangered species. Nest boxes are an effective conservation tool to improve productivity in areas lacking natural tree cavity nesting sites. Breeding success (calculated as the proportion of nesting attempts that fledged a chick) and predation levels were similar for groups using nest boxes and natural nests. Natural nests were more buffered against cooling night temperatures, but otherwise nest boxes provided nesting conditions that were no better than natural nests. Timing of breeding for nests in natural tree cavities and nest boxes were similar. However, groups with access to a nest box attempted breeding more often than groups with access to a natural nest only, resulting in an 15 % increase in the number of fledglings per group compared to an adjacent protected area with no artificial nests. The number of breeding groups in the study area increased by 460 % over 12 years. However, there is a limit to the density of breeding groups. Breeding success was highest when breeding density was one breeding group per 90-120 km², so nests should be spaced ~10 km apart. Given that the threats to ground hornbills include persecution and poisoning, increasing the reproductive rate by providing nest boxes should assist in slowing the decline by the increased recruitment of offspring into the population. Timing of breeding varied across years. The first eggs laid each year ranged from 9 September to 14 November, and median lay date was 03 November. Breeding attempts that were initiated early in the season were more likely to fledge a chick than those initiated later in the season. Timing of breeding was delayed during warmer springs, particularly under dry conditions. In savannas, hotter spring temperatures could limit food availability, for example, if higher temperatures cause the vegetation to dry out, resulting in a rapid decline in insect flush, especially in the phytophagous insect groups that form a large part of the ground hornbill diet. Factors to consider when constructing and placing nest boxes include thickness of the cavity walls, entrance height above ground and density of nest boxes placed in the landscape. Breeding attempts in natural nests and nest boxes with thicker nest walls and those positioned with higher entrances above the ground increased breeding success. Therefore, nests should be constructed with cavity walls at least 6 cm thick and placed so that the entrances are situated > 6 m above the ground. With 186 ringed chicks fledging from the study area after the installation of nest boxes, it was possible to observe their dispersal within the study area and farther away into the adjacent Kruger National Park. There was no evidence for sex-biased dispersal. Males and females dispersed at similar ages, and over similar distances, raising interesting questions about inbreeding avoidance mechanisms in this species. If females do not disperse beyond the range of related males, how do related individuals avoid pairing, and what forms of individual recognition exist?

Different histories for forests in Southern Africa have been hypothesized from vegetation biogeography and pollen analysis. However, the history of forests is still controversial. Phylogeography uses gene genealogies to infer history of distributions. Two forest endemic moss species were sampled: Leptodon smithii; and Neckera valentiniana. Two gene regions were used, trnLF (chloroplast genome) and ITS1 (nuclear genome). Neckera valentiniana showed no variation from the populations sampled. Results from Leptodon smithii based on the trn and ITS region suggest that forests once were widespread, but then became fragmented. Dispersal corridors still exist between Southern Cape populations and Western Cape populations.

The effective conservation of threatened and endangered plants requires an understanding of population dynamics and the evaluation of factors that could reduce population growth. I constructed and analyzed a stage structured demographic model for Euphorbia telephioides, a threatened dioecious perennial herb, to determine the current status of three populations, compare projections of population growth using different methods, and determine the effects of pollen limitation in the population dynamics of this species. Dioecious plants are prone to pollen limitation due to their inability to self-pollinate. Studies indicate that pollen limitation reduces seed set in plants due to insufficient quantity or quality of pollen, which can reduce population growth rate due to the decrease in fecundity. I combined experimental tests for pollen limitation with construction and analysis of structured demographic models, to examine how increased levels of pollen limitation would affect population growth rates. Determining the current status of populations, and simulating the consequences of possible threats, such as pollen limitation, provides a quantitative basis for conservation actions. I compared deterministic and stochastic projections of a stage structured demographic model to examine how environmental variation affects population growth rates, and I examined the effects of parameterizing the model excluding demographic measures of randomly marked individuals in the population growth rates (Lambda). The majority of estimated lambdas and their 95% confidence intervals indicate that these three populations are projected to decline. Lambda estimated excluding randomly marked individuals overestimated population growth because adult plants had 100% survival. I did not find evidence of significant pollen limitation of fruit or seed production, and simulations of increased levels of pollen limitation reduce Lambda at a modest rate between 0.17% to 1.91%. The main advantage of constructing a structured demographic model is that these models allow us to integrate data on different stages of a complex life cycle. In the case of E. telephioides elasticity analysis indicates that increasing stasis of non-flowering plants could lead to increasing population growth rates. / A Thesis submitted to the Department of Biological Science in partial fulfillment of the Master of Science. / Fall Semester 2016. / November 2, 2016. / Euphorbia telephioides, Population ecology / Includes bibliographical references. / Alice Winn, Professor Directing Thesis; Joseph Travis, Committee Member; Nora Underwood, Committee Member.

The environmental impacts of solar power generation and particularly Concentrated Solar Power (CSP) are not well understood. There have been reports of birds injured and killed by concentrated solar radiation at power 'towers' and from collisions with mirrors at both tower and 'trough' facilities. This study assesses the impacts of a utility-scale 50 MW 'trough' CSP facility - Bokpoort CSP Power Plant - in the Northern Cape, South Africa. To assess the changes in functional and structural changes in bird communities, bird counts in the solar fields (mirror arrays) were compared with transects from rangelands adjacent to the CSP plant. Invertebrates were sampled with sticky and pitfall traps adjacent to the power block, in the solar fields, and in the rangeland landscape to assess changes in invertebrate communities. There were significant changes in bird distribution across the landscape with more species richness and two orders of magnitude greater abundance in the rangeland compared to the solar fields. Fewer invertebrates were caught, but with a greater taxonomic richness in rangeland compared to the power block and solar field. The facilities' evaporation ponds created novel wetland habitat for birds; 23 species were recorded that would have been absent from the area prior to construction, including three breeding species. The solar fields were surveyed for bird injuries or fatalities over 3 months; only eight dead birds were found, all but one was too old to determine the cause of death (>1 month); the remaining carcass likely died from a mirror impact. Western Barn Owls (Tyto alba) made up half of the mortalities. Biases in mortality estimates due to searcher efficiency and scavenger removal were substantial only for small birds. Twenty-one animals (3 reptiles, 12 mammals, 6 birds) likely drowned in the evaporation ponds after being unable to escape. The recorded mortalities were very low in comparison with similar studies on CSP facilities. No threatened or endangered species were killed. Overall, the facility had a low impact on bird populations, but the drowning risk posed to animals by evaporation ponds requires mitigation. The negative impacts observed could be minimised through careful site selection of solar facilities and careful design and mitigation considerations particularly with regard to evaporation and water ponds in arid areas such as the Northern Cape.