Philosophiae Doctor - PhD / In the marine environment, penguins have been described as curators and serve a critical role in ecological balance. The African penguin (Spheniscus demersus) has undergone a rapid population decline, mainly due to disturbances in their natural habitat. The African penguin was up-listed from vulnerable to endangered on the IUCN Red List for Threatened Species in 2010 and thus urgent conservation action is required. Integral to long-term conservation action of any species is a basic knowledge of its reproductive biology, which is currently lacking for African penguins. The main aim of this investigation was to evaluate techniques for the collection of semen in African penguin and to determine sperm quality in order to cryopreserve sperm for in vitro fertilization (IVF) purposes of captive and wild populations. Semen was collected once a week during two breeding seasons from two captive African penguins. Ejaculates (n=51) were obtained over two breeding seasons (Jan-Feb and Jun-Oct) and evaluated for semen volume, sperm concentration, sperm vitality, sperm motility and sperm morphology. In addition twelve (six females and six males, n=4 were breeding pairs) captive African penguins were monitored for hormone (estradiol, testosterone, progesterone) levels prior to and after the egg-laying period.

Magister Scientiae (Biodiversity and Conservation Biology) - MSc (Biodiv and Cons Biol) / African penguin (Spheniscus demersus) mainland colonies are a popular tourist attraction in the Western Cape of South Africa. The African penguin population is in decline and the species is listed as endangered on the International Union for the Conservation of Nature (IUCN) Red List. This thesis aimed to investigate the impact of ecotourism upon African penguin colonies by comparing two colonies of differing levels of tourist visitation in the Western Cape in 2017. The high visitation colony is a mainland colony where ecotourism activities take place (Stony Point), and the low visitation colony is an island colony where ecotourism does not occur (Robben Island). As well as inter-colony comparisons, nests at the high visitation colony within areas of differing exposure levels were also compared.

Cape St Francis, Eastern Cape, has been identified as one of four potential sites for establishing a mainland-based African penguin (Spheniscus demersus) colony. This thesis comprises three main components: a verification of a preparation method for stable isotope samples from penguin feathers; a dietary analysis of the penguins on Bird Island, Algoa Bay, though stable isotope analysis of whole blood and feathers (2012 and 2013); and an estimation of available fish surplus that could potentially support a colony of penguins at Cape St Francis. Each component contributes towards the next, all building towards answering the main research question: Will there be enough food around St Francis Bay to support a colony of penguins and sustain the already established fisheries industry within the bay? Stable isotope analysis of whole blood and feathers from breeding adults and whole blood from juveniles provided insight into the variability of African penguins’ diets at different stages in their life history. Stable isotope mixing models indicated that the predicted proportions that each prey species could potentially contribute to diet conflicted with published stomach sample data. This might arise from inaccurate trophic enrichment factors used in the model, or from systematic biases in the published stomach sampling techniques, or both. Dietary sexual dimorphism was not demonstrated by the isotope signatures of breeding penguins. Based on official catch data, the fisheries activity on the south coast, and especially around the potential colony site at St Francis, is much lower than around the west coast’s penguin colonies. The model provided a first-order estimate for fish supply around the potential colony site at St Francis both at a large coastal scale and a local small scale. At both scales the estimate indicated an ample availability of fish at current fishing levels. The model in Chapter 4 can also be applied to refining the assessments of other potential colony sites on the south coast. In conclusion, the south coast is a promising area for a new colony of penguins in terms of food availability. There is relatively low fishing activity in the area and, as suggested by the large-scale model in Chapter 4, an ample fish resource. The final chapter briefly discusses factors that need to be considered before attempting to establish a mainland-based colony of African penguins.

Stable isotope (SI) ratios of carbon (δ13C) and nitrogen (δ15N) are now widely used as biomarkers in ecological studies to provide information about food web structuring. However, understanding trophic relationships using SI analysis requires not only knowledge of SI values of predator and prey, but also accurate discrimination factors (DFs), which can differ among species and by physiological state. This thesis examined three questions using captive birds from the South African Foundation for the Conservation of Coastal Birds (SANCCOB). First, the effects of ontogeny on δ13C and δ15N ratios of African penguins (Spheniscus demersus) were assessed. Blood samples were collected from penguins in four age classes (P3 chicks, blues, juveniles and adults) concurrently with their diet (sardine (Sardinops sagax) and formula). Second, to assess the influence of breeding physiology on SI ratios, the blood of ten breeding pairs of penguins was sampled over a five-month period from June to October 2016. Following laying, each pair was categorised into one of three (four for whole blood) egg production phases (initial yolk deposition, rapid yolk deposition and post-laying) and their influences on SI ratios were tested. Third, species differences in DFs were evaluated for African penguins, kelp and Hartlaub’s gulls (Larus dominicanus and L. hartlaubii), greater crested terns (Thalasseus bergii) and Cape cormorants (Phalacrocorax capensis). Flying birds were mostly fed sardine with a small but unknown amount of sardinella (Sardinella aurita), DFs were therefore estimated for a 50:50 sardine:sardinella diet, a 75:25 sardine:sardinella diet and a 100% sardine diet for each flying bird species. The DFs were assessed for the whole blood (WB), red blood cells (RBC), plasma (PL) and delipidated plasma of the penguins, and only WB for the flying birds as well as flesh, whole fish, delipidated flesh and delipidated whole fish for fish species, and for formula. Results indicated that age influenced both the δ13C and δ15N of WB, only the δ15N of RBC and the δ13C of delipidated PL. The assessment of breeding physiology yielded a significant interaction between the effects of egg production phase and sex on the δ13C of WB; females had significantly lower δ13C in the rapid yolk deposition phase than the other two phases and all males. The δ13C of PL was affected only by sex, with females having a significantly lower δ13C value than males. Neither physiological state nor sex influenced the other blood components. Differences were found among the three DFs in the non-penguin species, but not for all consumer – prey tissue combinations. There were also significant differences among species with a DF calculated from a diet with the most probable prey proportions eaten. Depending on the combination of consumer and prey tissue used to calculate the DF, a different conclusion regarding trophic information can be reached. A literature review updated with the present data showed that no general pattern or grouping of similar species with regards to DF values could be drawn, highlighting the importance of determining species- and tissue-specific DFs. Thus age, egg production, tissue and species all influenced the SI values of bird blood and therefore their DFs. Not all physiological conditions affect all blood components in the same way, making different components more or less sensitive to physiological influences. Though their influence is at a small enough scale that it is unlikely to hamper correct conclusion in ecological studies, it is crucial that these factors are considered when using SI analysis (SIA). When uncertainties exist for some coefficients in wild studies, SIA should therefore be combined to other dietary techniques to determine the food web structure as best as possible.

Animal behaviour is especially sensitive to environmental variability and prey availability during the breeding season, and this is particularly true for non-volant, central place foragers such as the endangered African Penguin (Spheniscus demersus). Individual sex and morphology, as well as the level of assortment within mated pairs can influence both the behaviour and the reproductive success of species exhibiting biparental care. This study made use of a large biometric database and nest attendance video footage to determine the influence of intrinsic (assortative mating, brood size and chick age) and extrinsic (environmental conditions, anthropic disturbance) factors on breeding behaviour and performance of African Penguins on Bird Island, Algoa Bay, during peak breeding (March - July) in 2013. While sexual dimorphism in African Penguins is subtle, the colony-specific discriminant functions presented here provide an accurate sexing tool when only bill and flipper length are available. Despite the premise that selection of a large, high-quality mate in this longlived, monogamous seabird governs lifetime fitness, only low levels of assortative mating were found, and this only for earlier breeders, when larger females (but not males) bred. The 2013 season was a particularly successful one, coinciding with above-average sardine and anchovy abundance, and almost 80 percent of monitored nests were double-brooded, with very low levels of mortality. A- and B-chicks of double broods and singleton chicks grew at similar rates and exhibited similar body condition indices. In these conditions, chick developmental rates were independent of parental size, assortment or provisioning behaviour. Females raising a double brood were significantly lighter and in poorer body condition than those raising a single chick, although the same trend was not evident in males. Offspring sex ratio in 2013 (2.27:1) favoured male chicks, suggesting that there is potential to over-produce the larger sex when resources are plentiful. Peak nest arrival and departure times of parents did not change over the course of monitored breeding attempts (March-June), nor were they different for disturbed and undisturbed nests or for a single or double brood. The increase in CCTV-observed provisioning rate as chicks grew larger was best explained by brood size, at-sea chlorophyll a concentration, and maximum air temperature, but was unrelated to parental morphology or assortative index. Importantly, parental absenteeism commenced earlier and was markedly greater in nests frequently handled by researchers than in undisturbed nests. Both the time spent together by parents, and absenteeism were measurably affected by maximum afternoon air temperatures, the effects of which are expected to be exacerbated by poorer foraging conditions and climate change. A third of manually-monitored nests shared chick-guarding duties unequally, although this phenomenon was independent of parental sex or morphology. The adaptive benefits of mating patterns and division of labour during chick-rearing may only become apparent in a year of below-average food availability and it is highly recommended that this study be repeated in a year of scarce food resources. These findings augment past foraging ecology studies and demonstrate that investigator disturbance and environmental conditions can affect the nesting behaviour of this highly threatened seabird.

This study investigated the potential impacts of several anthropogenic stressors on the movement and survival of African penguins in Algoa Bay, taking into account prey availability and environmental conditions (sea surface temperature and chlorophyll productivity). The primary drive for the study was based on the seismic exploration, which took place in the habitat of the endangered African penguin. The research was also carried out to understand how pelagic fishing and prey availability influence the broader behavioural and mortality aspects of penguins. To achieve this goal, I used monthly beach surveys to assess seabird mortality, admissions of penguins in rehabilitation centres and the monitoring of breeding penguins’ movement at sea through individual GPS tracking. In addition, prey distribution and abundance were monitored with acoustic surveys to assess potential changes in relation anthropogenic stressors. Breeding African penguins did not react dramatically to seismic activities, except on St Croix Island where they changed their foraging direction during the seismic period. There was no evidence of a siginficant increase in African penguin carcasses encountered or strandings along the beaches during the period of seismic activity. However, a sharp increase in encounter rate was recorded soon after seismic operations, which may potentially indicate a delay between the effect of seismic activities on penguin mortality and the encounter of the related carcasses after the operation. However, the impact of seismic surveys on penguins may be long-term rather than short term, and additional studies would be needed to reveal long-term effects of seismic activities, if any. Foraging effort of breeding African penguins increased substantially in the presence of commercial fishing activities within close proximity to the colonies. To a greater extent, at-sea movement patterns and counts of beach cast carcasses were largely influenced by prey availability. Penguins spent more time at sea, traveled longer and covered a larger foraging area during periods of relatvely low pelagic fish abundance. There was also a sharp increase in African penguin mortality during periods of low prey availability in the Bay. However, environmental conditions showed some influence over these interactions. Foraging effort of breeding African penguins increased substantially in the presence of commercial fishing activities within close proximity to the colonies. To a greater extent, at-sea movement patterns and counts of beach cast carcasses were largely influenced by prey availability. Penguins spent more time at sea, traveled longer and covered a larger foraging area during periods of relatvely low pelagic fish abundance. There was also a sharp increase in African penguin mortality during periods of low prey availability in the Bay. However, environmental conditions showed some influence over these interactions. For example a red tide event during in period of low prey availability, made it difficult to understand impacts of prey on penguins difficult. Both anthropogenic stressors and environmental conditions influenced African penguins’ behaviour and survival. Due to the worrying trend of African penguins in South Africa for the past few years, all conservation management efforts to increase penguin numbers and limit their mortality are necessary. This study highlights the negative impact of industrial fishing on this Endangered species and the rapid increase in mortality of penguins during times of low prey availability. A network of Marine Protected Areas would certainly increase food availability to African penguins by limiting competition with fisheries, and contribute to increasing the population numbers.