The distribution and subsequent availability of marine predators’ prey is highly variable and is linked to fluctuating oceanographic parameters. It is well documented that annual breeding success of seabirds is related to the temporal availability of prey. Knowledge of a seabird’s diet and at sea distribution is therefore critical in deducing the effect of a fluctuating environment on a seabirds’ population and, furthermore, to understand seabirds role in the environment. The gentoo penguin population at sub-Antarctic Marion Island experienced a decline of 52% between 1994 and 2012. It has been speculated that long term changes in the local marine environment, due to a southward shift of the sub-Antarctic Front (SAF), and subsequent changes in prey availability is the ultimate causal factor. Therefore, this thesis investigates the temporal variation in the diet these birds over three years, through the use of stomach content samples. It then investigates whether the composition of the diet has changed over the long term (approximately 18 years). Additionally, the at sea distribution of these birds was investigated using temperature-depth recorders (TDRs) and global positioning system (GPS) devices. This study represents the first to investigate variation in the diet over three years and over the long term. It is also the first study to track these birds fine scale movement. Over three years, the diet exhibited a high degree of variability. The birds primarily foraged on the fish, Lepidonotothen larseni and the crustaceans, Nauticaris marionis and Euphausia vallentiniy. Over three years, L. larseni became increasingly more important during the breeding season. This was attributed to an increase in availability of these fish during this time, as this time corresponded to the transition of these fish from the larval pelagic phase to the benthic adult phase. Furthermore, the proportions of N. marionis and E. vallentini in the diet varied significantly among years. This variation is possibly a result of the highly variable latitudinal position of the SAF. The diet changed over the long term. There was significantly less N. marionis and more E. vallentini in the diet during more recent years. The long term southward shift of the SAF is considered to be a leading cause of these significant changes in the diet. The shallow inter-island shelf between Marion and Prince Edward Island was an important foraging area possibly due to predictable prey resources. In addition, these birds exhibited an important and possibly unique foraging strategy. They performed short self-maintenance trips where after they return to land to roost for the night. The following day these trips were followed by a longer foraging trip that was followed by chick provisioning. This strategy is unique among seabirds longer trips are associated with self-maintenance and not chick provisioning. This study has provided important knowledge of the foraging ecology of the gentoo penguin at sub-Antarctic Marion Island. It highlights the necessity for continuation of diet and tracking studies of these birds if we are to fully understand the ultimate factors that are causing the decline in their breeding success and population numbers.

Marine top predators often occupy broad geographical ranges that encompass varied habitats. Therefore, a pre-requisite towards conserving these animals is to determine the components of their realized niche, and investigate whether a species is a specialist or a generalist. For generalist species, it is also necessary to understand if local specialisation occurs. Uncovering these components can allow us to build models of a species realized niche that may then be used to infer habitat use in unsampled locations. However, fully understanding the components of a marine top predators realized niche is challenging owing to the limited opportunity for in situ observations. Overcoming these limitations is a key step in marine top predator research. It will enhance our understanding of trophic coupling in marine systems, and aid in the development of tools to better study these predators in their dynamic environment. Seabirds, penguins (Spheniscids) in particular, are a group of animals for which investigating their realized niche is of vital importance. This is because numerous species face growing uncertainty in the Anthropocene, and in a time of rapid environmental change there is furthermore a need to better understand the potential use of these birds as indicators of ecosystem health. The aim of this thesis, therefore, is to investigate the foraging ecology of gentoo penguins (Pygoscelis papua) at the Falkland Islands. At the Falkland Islands, limited historical information exists regarding this species foraging ecology, with most information coming from a single location at the Falklands. As the Falkland Islands have the world’s largest population of gentoo penguins, elucidating factors influencing this population will have global relevance. Furthermore, historical information indicated potential competition with fisheries, and with prospecting for hydrocarbons and an inshore fishery, there is a need to understand the distribution of these birds across the islands. Penguins are also well suited to carry biologging devices allowing for in situ observations of inter and intraspecific interactions, as well as habitat specific interactions. In this study, I sampled birds over three breeding seasons, from four breeding colonies - chosen for their varied surrounding at sea habitat - across the Falkland Islands. I investigated the diet with stomach content and stable isotope analysis, the at-sea distribution with GPS and time depth recorders, and how these birds behaved at sea using custom made animal-borne camera loggers. Furthermore, I developed a method to recognise prey encounter events from back mounted accelerometers, using a supervised machine learning approach. As part of the first species specific description of diet at this scale for the Falklands, I revealed six key prey items for the birds: rock cod (Patagonotothen spp.), lobster krill (Munida spp.), Falkland herring (Sprattus fuegensis), Patagonian squid (Doryteuthis gahi), juvenile fish (likely all nototheniids), and southern blue whiting (Micromesistius australis). The use of animal-borne camera loggers verified that not only do gentoo penguins consume a diverse array of prey items, but they adopted various methods to capture and pursue prey, with evidence of birds following optimal foraging theory. Prey composition varied significantly between study sites with the at-sea distribution and habitat use of penguins reflecting that of local prey. Birds from colonies close to gently sloping, shallow waters, foraged primarily in a benthic manner and had larger niche widths. However, those at a colony surrounded by steeply sloping, deeper waters, typically foraged in a pelagic manner. Contrasting diet patterns were also prevalent from stable isotope data, and the niche widths of birds relating to both stomach content and stable isotope data were larger at colonies where benthic foraging was prevalent. Therefore, it was clear that surrounding bathymetry played a key role in shaping this species’ foraging ecology, and that at the population level at the Falkland Islands birds are generalists. However, at individual colonies some specialisation occurs to take advantage of locally available prey. I developed habitat distribution models - via boosted regression trees – which transferred well in time but poorly across space. Reasons for poor model transfer might relate to the generalist foraging nature of these birds and the reduced availability of environmental predictors owing to the limited range of these birds. I furthermore developed a method to identify prey encounter events that can also, to a degree, distinguish between prey items. This method will be a promising approach to refine habitat distribution models in future. These habitat distribution models could potentially contribute to marine spatial planning at the Falkland Islands. Footage from animal-borne camera loggers clearly showed that prey behaviour can significantly influence trophic coupling in marine systems and should be accounted for in studies using marine top predators as samplers of mid to lower trophic level species. Ultimately, flexibility in foraging strategies and inter-colony variation will play a critical role when assessing factors such as interspecific competition or overlap with anthropogenic activities.

Knowledge of diet and distribution in marine top predators is critical in interpreting their ecological role. Furthermore, this information is required for effective management and conservation of these species. For gentoo penguins at the Falkland Islands a previous overlap in dietary items with commercial fisheries was recognised. Despite this the at-sea distribution has been poorly studied and only at a few localities. Therefore, this thesis looks to broaden the knowledge of foraging ecology for gentoo penguins at another site in the archipelago, Cow Bay. The aim was firstly, through stomach content analysis, to determine if diet changed over a large temporal scale (2002/03/04 – 2011/12/13), and between the chick guard and crèche periods of chick rearing. Secondly, through GPS loggers, to assess distribution at sea during the incubation and chick guard periods (2012/13). Additionally, foraging ecology between genders was investigated. Gentoo penguins at Cow Bay foraged primarily on fish (66-99 percent) throughout the entire study period. The principle prey item by mass was Patagonotothen spp., considered previously as discard by fisheries but now an important commercial resource, for all periods (47-78 percent) except that of the 2012/13 crèche period (19 percent). During this period Sprattus fugensis made up the bulk of the diet (52 percent). Cephalopods, particularly Loligo gahi (1-24 percent), were also prominent in the diet and crustaceans contributed negligibly. The diet indicated that gentoo penguins are capable of both a benthic and pelagic feeding strategy which likely reflects changes in local prey abundance. Birds typically conducted foraging activities during daylight hours with greater trip duration, path length, maximum distance and area utilised during the incubation period than the guard. All trips were confined to the continental shelf and within the boundaries of a marine management zone, the Falkland Islands Interim Conservation Zone (FICZ). Males and females fed on the same size and composition of prey items. Trip duration, path length and maximum distance from the colony were not significantly different, although males foraged in a slightly larger area than females marginally to the north. As at-sea distribution was confined within the FICZ this management area provides protection to food resources of gentoo penguins breeding at Cow Bay. At this site, the population is increasing, likely associated with increased prey availability as evidenced by record catches of Patagonotothen spp. in recent years at the Falkland Islands. Continued long term monitoring of gentoo penguin diet and fishery yields to allow detection of possible competition in the future is recommended. As marine sentinels, on-going monitoring of gentoo penguin populations at the Falkland Islands is likely to provide early warning signs of reduced prey availability.

Submitted by William Justo Figueiro (williamjf) on 2015-08-11T21:04:47Z No. of bitstreams: 1 13e.pdf: 3156102 bytes, checksum: da00a19d9782f6ac017289a5d272ac1f (MD5) / Made available in DSpace on 2015-08-11T21:04:47Z (GMT). No. of bitstreams: 1 13e.pdf: 3156102 bytes, checksum: da00a19d9782f6ac017289a5d272ac1f (MD5) Previous issue date: 2013 / CAPES - Coordenação de Aperfeiçoamento de Pessoal de Nível Superior / CNPQ – Conselho Nacional de Desenvolvimento Científico e Tecnológico / INCT-APA - Instituto Nacional de Ciência e Tecnologia Antártico de Pesquisas Ambientais / PROANTAR - Programa Antártico Brasileiro / SECIRM - Secretaria Interministerial para os Recursos do Mar / MMA - Ministério do meio Ambiente / INCT-APA - Instituto Nacional de Ciência e Tecnologia Antártico de Pesquisas Ambientais / UNISINOS - Universidade do Vale do Rio dos Sinos / O nicho realizado de duas espécies precisa ser diferente, para que estas espécies possam coexistir de maneira estável. Pygoscelis papua e P. antarcticus reproduzem simpatricamente na Ilha Elefante, Antártica. Foram coletadas amostras de conteúdo estomacal e realizado biometria das duas espécies para analisar a dieta e a sobreposição de nicho. Um total de 56 amostras de P. papua e 71 amostras de P. antarcticus foram coletadas durante dois períodos de reprodução austral, 2010/11 e 2011/12, na região de Stinker Point. A proporção de itens variou entre as espécies de pinguins e entre os táxons representados. E. superba foi a presa de maior abundância comparado com outros itens alimentares, com aproximadamente 69% FO para P. papua e 98% FO para P. antarcticus. Foram encontradas nove espécies de peixe, seis espécies de crustáceos e uma espécie de cefalópode, identificados a nível específico. Existe uma diferença significativa entre os fatores, espécie de pinguim, tamanho da carapaça e sexo de krill antártico, quando comparados entre si. Além disso, a biometria indica que o comprimento do bico e peso de P. papua são maiores que em P. antarcticus, da mesma forma que o bico é mais alto e mais largo em P. antarcticus. Tais diferenças morfológicas podem explicar as diferenças no forrageio. Foi observada uma sobreposição de nicho trófico entre as espécies, pela utilização dos mesmos recursos, uma vez que estas espécies ocorrem simpatricamente na mesma região. Este estudo demonstra que a variação específica do nicho trófico ocupado pelas espécies pode ser definida pelo comportamento de forrageio e pela seleção dos recursos alimentares de cada espécie. É importante a recomendação destas espécies como indicadores de qualidade ambiental, adicionando questões como a variabilidade local, pois o nicho trófico pode alterar ao longo do tempo. / The two species realized niche needs to be different, since these species can coexist stably. Pygoscelis papua and P. antarcticus breed simpatricaly on Elephant Island, Antarctica. Stomach content samples were collected and biometrics of both species was measured in order to analyze the diet and the niche overlap. A total of 56 P. papua samples and 71 P. antarcticuss samples were collected, during the two austral breeding seasons, 2010/11 and 2011/12, on the Stinker Point region. The proportion of items ranged from the penguin species and among the taxa represented. E. superba was the most abundant prey compared with other food items, with approximately 69% FO for P. papua, and 98% FO for P. antarcticus. We found nine species of fish, six species of crustaceans and one species of cephalopod, identified by the specific level. There is a significant difference between the factors, species of penguin, carapace length and sex of antarctic krill, when compared with each other. In addition, the biometrics indicates that length of P. papua’s beak and weight are larger than P. antarcticus, but P. antarcticus has the highest and widest beak measures. Such morphological differences may explain differences in foraging. We observed a niche overlap of species, by the use of the same food resources, once these species occurs simpatricaly in the same region. This study also demonstrated that the specific variation of trophic niches occupied by the species may be defined by the foraging behavior and by the selection of the food resources. It is important the recommendation of these species as indicators of environmental quality, adding issues such as local variability since the trophic niche may change over the time.