Esta tese aborda a definição de um projeto estratégico-científico brasileiro para a Antártida, a partir da análise da evolução da geopolítica do Sistema do Tratado Antártico (STA). A questão antártica, conforme esta tese, se fundamenta em dois pressupostos teóricos antagônicos das relações internacionais, o realismo e o liberalismo. As premissas de cooperação científica e de uso pacífico da região, estabelecidos no Tratado da Antártida (1959), determinaram a discussão do objeto de estudo a partir de uma epistemologia liberal. A geopolítica antártica, neste início do século XXI, pende para uma dialética científico-ambiental, em detrimento da dimensão econômico–territorialista que dominou o discurso para a região austral até o emblemático ano de 1991, quando ocorreu a ratificação do Protocolo sobre Proteção Ambiental do Tratado da Antártida (Protocolo de Madri). Se o século XX foi marcado pelo advento da ciência antártica, representado pelo Ano Geofísico Internacional (1957 - 1958), o presente século se caracteriza, até o momento, como aquele em que essa ciência antártica deverá construir sua própria agenda. Os Estados-signatários, cada vez mais, reconhecem o peso político de uma ciência antártica de excelência dentro do STA. Com a instituição do Programa Antártico Brasileiro (PROANTAR), em 1982, o país passou a defender seus substanciais interesses naquela região, que, em um primeiro momento, estavam subordinados às contingências da Guerra Fria e ao Projeto do Brasil-potência. Todavia, a invocação subjetiva de tais interesses não permitiu que se delimitasse a sua real amplitude. Esse posicionamento geopolítico em relação à Antártida começou a ser referenciado a partir de 2011, quando a questão antártica, através da elaboração de um Planejamento Estratégico para o PROANTAR (2012 - 2022), passou a assumir uma relativa importância estratégica para o país; em outras palavras, o programa aproximou-se de um projeto de Estado. No referido planejamento encontram-se as diretrizes que deverão garantir ao Brasil um status político relevante dentro do STA, o que implica numa participação efetiva do país nas decisões sobre o destino daquela região. A diretriz principal desse processo é a inserção do país no seleto grupo de Estados-signatários que desenvolve pesquisas de ponta na Antártida. Ao Entender a importância estratégico-geopolítica da Antártida para o Brasil, e o protagonismo da ciência na política do STA, esta tese propõe novos desafios e paradigmas à ciência antártica brasileira, apoiando o atual Planejamento Estratégico para o PROANTAR (2012 - 2022) e o mais recente Plano de Ação para a Ciência Antártica (2013 - 2018). Assim, analisar o PROANTAR a partir da definição de um projeto estratégico-científico, e suas repercussões geopolíticas, é o objetivo maior desta tese, implicando nas seguintes metas: uma análise sobre a geopolítica antártica, o que determinou uma discussão preliminar sobre os pressupostos geopolíticos do Tratado e da ciência antártica, a partir de uma perspectiva histórica; uma análise sobre as dimensões da geopolítica da região no limiar do século XXI, identificando o ano de 1991 (ratificação do Protocolo de Madri) como o momento de inflexão do STA, no qual a dimensão científico-ambiental começou a adquirir maior relevância geopolítica; uma análise sobre a questão antártica no pensamento geopolítico brasileiro, que permitiu discorrer sobre a construção de um possível imaginário territorialista antártico, assim como a sua repercussão na dimensão científica do PROANTAR. Por fim, se analisa os novos desafios à ciência antártica brasileira no século XXI: a expansão geográfica das pesquisas do PROANTAR naquele continente, em especial as investigações no interior do manto de gelo antártico, que quebra o paradigma de uma ciência periférica, historicamente restrita à região da Península Antártica; e a instituição de uma política capaz de garantir a qualidade da produção científica, através de um adequado apoio logístico-financeiro. Vencer tais desafios é de importância fundamental para a inclusão da pesquisa antártica brasileira nas chamadas fronteiras emergentes da ciência. Todavia, à medida que se analisa as ações político-científicas, convergentes à definição de um projeto estratégico brasileiro para a Antártida, que vêm sendo implementadas desde o início deste século, mais nítida se tornam as contradições e os obstáculos à definição de tal projeto: os entraves burocráticos e a volatilidade das ações políticas, que, somados a falta de reconhecimento da importância (geo)política de uma ciência antártica de vanguarda, em particular por grande parte dos pesquisadores que atuam no PROANTAR, impedem a instituição de uma política científica e a definição de um projeto estratégico-científico para a região. / This thesis addresses the definition of a Brazilian strategic-scientific project for Antarctica from the analysis of the Antarctic Treaty System (ATS) geopolitical evolution. The Antarctic question, as discussed in this thesis, is based in two antagonistic theoretical premises taken from the foreign relations (realism and liberalism). The premises of scientific collaboration and the pacific use of the region, established in the Antarctic Treaty (1959), determined a discussion of this topic using a liberal epistemology. The Antarctic geopolitics at the beginning of the 21st century leans towards a scientific-environmental dialectics, to the detriment of the economical-territorial dimension that dominated the discourse for the Austral region up to the emblematic year of 1991, when the Protocol on Environmental Protection to the Antarctic Treaty (Madrid Protocol) was ratified. If the 20th century was marked by the arrival of the Antarctic science, represented by the International Geophysical Year (1957-1958), the current century is characterized, so far, as the one in which the Antarctic science should build its own agenda. The Signatory-States increasingly recognize the political weight of an Antarctic science of excellence within the ATS. By establishing the Brazilian Antarctic Program (PROANTAR, from the Portuguese Programa Antártico Brasileiro) in 1982, Brazil started to defend of its substantial interests in the Antarctic region. These interests, at first, were subordinated to the Cold War contingencies and the Brazil-potency Project. However, the subjective invocation of such interests did not allow the delimitation of its real extent. This geopolitical positioning in relation to Antarctica was first mentioned in 2011 when the Antarctic question, through the preparation of a Strategic Planning for PROANTAR (2012 - 2022), acquired a relative strategic importance for the country. In other words, the program became closer to a project of State. This plan contains directives that shall grant a relevant political status to Brazil within the ATS, which implies the effective participation of the country in the decisions about the fate of the region. The integration of the country into the select group of signatory States that develop cutting-edge research in Antarctica is the main directive of this plan. By understanding the strategic-geopolitical importance of Antarctica to Brazil, and the role of science in the ATS politics, this thesis proposes new challenges and paradigms to the Brazilian Antarctic science, supporting to the current Strategic Planning for PROANTAR (2012 - 2022) and the most recent Action Plan for the Antarctic Science (2013 - 2018). Thus, the analysis of PROANTAR from the perspective of the definition of a strategic-scientific project and its geopolitical repercussions are the main objetives of this thesis, which are reflected in the following goals: an analysis of the Antarctic geopolitics, which determined a preliminary discussion regarding the geopolitical premises of the Treaty and the Antarctic science from a historical perspective; an analysis of the geopolitical dimensions of the region at the benning of the 21st century, identifying the year 1991 (ratification of the Madrid Protocol) as the ATS turning point, when the scientific-environmental dimension began to acquired a greater geopolitical relevance; an analysis of the Antarctic question in the Brazilian geopolitical line of thought, which allowed the discussion about the construction of a possible Antarctic territorial imaginary, as well as its repercussion on the scientific dimension of PROANTAR. Finally, we analyze the new challenges to the Brazilian Antarctic science in the 21st century: the geographic expansion of the PROANTAR research in that continent, especially the investigations into the Antarctic ice sheet, breaking the paradigm of a peripheral science that has been historically confined to the Antarctic Peninsula region; and the institution of a policy that guarantees the quality of the scientific production through an adequate logistic-financial support. Overcoming such challenges is paramount for the inclusion of the Brazilian Antarctic research in the so-called emerging frontiers of science. Nonetheless, as we analyze the political-scientific actions, converging to the definition of a Brazilian strategic project for Antarctic, which has been implemented since the beginning of the century, it becomes clearer the contradictions and obstacles to the definition of such project become: the bureaucratic hindrances and the volatile political actions, which in addition to the lack of recognition of the (geo)political relevance of a cutting-edge Antarctic science, especially by a large part of the scientific community working at PROANTAR, prevent the institution of a the scientific politicy and the definition of a strategic-scientific project for the region.

<p align="justify">The Southern Ocean (ca. 20% of the world ocean surface) is a key place for the regulation of Earth climate thanks to its capacity to absorb atmospheric carbon dioxide (CO2) by physico-chemical and biological mechanisms. The biological carbon pump is a major pathway of absorption of CO2 through which the CO2 incorporated into autotrophic microorganisms in surface waters is transferred to deep waters. This process is influenced by the extent of the primary production and by the intensity of the remineralization of organic matter along the water column. So, the annual cycle of sea ice, through its in situ production and remineralization processes but also, through the release of microorganisms, organic and inorganic nutrients (in particular iron)into the ocean has an impact on the carbon cycle of the Southern Ocean, notably by promoting the initiation of phytoplanktonic blooms at time of ice melting.</p><p><p align="justify">The present work focussed on the distribution of organic matter (OM) and its interactions with the microbial network (algae, bacteria and protozoa) in sea ice and ocean, with a special attention to the factors which regulate the biological carbon pump of the Southern Ocean. This thesis gathers data collected from a) late winter to summer in the Western Pacific sector, Western Weddell Sea and Bellingshausen Sea during three sea ice cruises ARISE, ISPOL-drifting station and SIMBA-drifting station and b) summer in the Sub-Antarctic and Polar Front Zone during the oceanographic cruise SAZ-Sense.</p><p><p align="justify">The sea ice covers were typical of first-year pack ice with thickness ranging between 0.3 and 1.2 m, and composed of granular and columnar ice. Sea ice temperature ranging between -8.9°C and -0.4°C, brines volume ranging between 2.9 to 28.2% and brines salinity from 10 to >100 were observed. These extreme physicochemical factors experienced by the microorganisms trapped into the semi-solid sea ice matrix therefore constitute an extreme change as compared to the open ocean. Sea ice algae were mainly composed of diatoms but autotrophic flagellates (such as dinoflagellates or Phaeocystis sp.) were also typically found in surface ice layers. Maximal algal biomass was usually observed in the bottom ice layers except during SIMBA where the maxima was localised in the top ice layers likely because of the snow and ice thickness which limit the light available in the ice cover. During early spring, the algal growth was controlled by the space availability (i.e. brine volume) while in spring/summer (ISPOL, SIMBA) the major nutrients availability inside sea ice may have controlled algal growth. At all seasons, high concentrations of dissolved and particulate organic matter were measured in sea ice as compared to the water column. Dissolved monomers (saccharides and amino acids) were accumulated in sea ice, in particular in winter. During spring and summer, polysaccharides constitute the main fraction of the dissolved saccharides pool. High concentrations of transparent exopolymeric particles (TEP), mainly constituted with saccharides, were present and their gel properties greatly influence the internal habitat of sea ice, by retaining the nutrients and by preventing the protozoa grazing pressure, inducing therefore an algal accumulation. The composition as well as the vertical distribution of OM in sea ice was linked to sea ice algae.</p><p><p align="justify">Besides, the distribution of microorganisms and organic compounds in the sea ice was also greatly influenced by the thermodynamics of the sea ice cover, as evidenced during a melting period for ISPOL and during a floodfreeze cycle for SIMBA. The bacteria distribution in the sea ice was not correlated with those of algae and organic matter. Indeed, the utilization of the accumulated organic matter by bacteria seemed to be limited by an external factor such as temperature, salinity or toxins rather than by the nature of the organic substrates, which are partly composed of labile monomeric saccharides. Thus the disconnection of the microbial loop leading to the OM accumulation was highlighted in sea ice.</p><p><p align="justify">In addition the biofilm formed by TEP was also involved in the retention of cells and other compounds(DOM, POM, and inorganic nutrients such as phosphate and iron) to the brine channels walls and thus in the timing of release of ice constituents when ice melts. The sequence of release in marginal ice zone, as studied in a microcosm experiments realized in controlled and trace-metal clean conditions, was likely favourable to the development of blooms in the marginal ice zone. Moreover microorganisms derived from sea ice (mainly <10 µm) seems able to thrive and grow in the water column as also the supply of organic nutrients and Fe seems to benefit to the pelagic microbial community.</p><p><p align="justify">Finally, the influence of the remineralization of organic matter by heterotrophic bacterioplankton on carbon export and biological carbon pump efficiency was investigated in the epipelagic (0-100 m) and mesopelagic(100-700 m) zones during the summer in the sub-Antarctic and Polar Front zones (SAZ and PFZ) of the Australian sector (Southern Ocean). Opposite to sea ice, bacterial biomass and activities followed Chl a and organic matter distributions. Bacterial abundance, biomass and activities drastically decreased below depths of 100-200 m. Nevertheless, depth-integrated rates through the thickness of the different water masses showed that the mesopelagic contribution of bacteria represents a non-negligible fraction, in particular in a diatom-dominated system./</p><p><br><p><p align="justify">L’océan Antarctique (± 20% de la surface totale des océans) est un endroit essentiel pour la régulation du climat de notre planète grâce à sa capacité d’absorber le dioxyde de carbone (CO2) atmosphérique par des mécanismes physico-chimique et biologique. La pompe biologique à carbone est un processus majeur de fixation de CO2 par les organismes autotrophes à la surface de l’océan et de transfert de carbone organique vers le fond de l’océan. Ce processus est influencé par l’importance de la production primaire ainsi que par l’intensité de la reminéralisation de la matière organique dans la colonne d’eau. Ainsi, le cycle annuel de la glace via sa production/reminéralisation in situ mais aussi via l’ensemencement de l’océan avec des microorganismes et des nutriments organiques et inorganiques (en particulier le fer) a un impact sur le cycle du carbone dans l’Océan Antarctique, notamment en favorisant l’initiation d’efflorescences phytoplanctoniques dans la zone marginale de glace.</p><p><p align="justify">Plus précisément, nous avons étudié les interactions entre le réseau microbien (algues, bactéries et protozoaires) et la matière organique dans le but d’évaluer leurs impacts potentiels sur la pompe biologique de carbone dans l’Océan Austral. Deux écosystèmes différents ont été étudiés :la glace de mer et le milieu océanique grâce à des échantillons prélevés lors des campagnes de glace ARISE, ISPOL et SIMBA et lors de la campagne océanographique SAZ-Sense, couvrant une période allant de la fin de l’hiver à l’été.</p><p><p align="justify">La glace de mer est un environnement très particulier dans lequel les microorganismes planctoniques se trouvent piégés lors de la formation de la banquise et dans lesquels ils subissent des conditions extrêmes de température et de salinité, notamment. Les banquises en océan ouvert étudiées (0,3 à 1,2 m d’épaisseur, températures de -8.9°C à -0.4°C, volumes relatifs de saumure de 2.9 à 28.2% et salinités de saumures entre 10 et jusque >100) étaient composées de glace columnaire et granulaire. Les algues de glace étaient principalement des diatomées mais des flagellés autotrophes (tels que des dinoflagellés ou Phaeocystis sp.) ont été typiquement observés dans les couches de glace de surface. Les biomasses algales maximales se trouvaient généralement dans la couche de glace de fond sauf à SIMBA où les maxima se trouvaient en surface, probablement en raison de l’épaisseur des couches de neige et de glace, limitant la lumière disponible dans la colonne de glace. Au début du printemps, la croissance algale était contrôlée par l’espace disponible (càd le volume des saumures) tandis qu’au printemps/été, la disponibilité en nutriments majeurs a pu la contrôler. A toutes les saisons, des concentrations élevées en matière organique (MO) dissoute et particulaire on été mesurées dans la glace de mer par rapport à l’océan. Des monomères dissous (sucres et acides aminés) étaient accumulés dans la glace, surtout en hiver. Au printemps et été, les polysaccharides dissous dominaient le réservoir de sucres. La MO était présente sous forme de TEP qui par leurs propriétés de gel modifie l’habitat interne de la glace. Ce biofilm retient les nutriments et gêne le mouvement des microorganismes. La composition et la distribution de la MO dans la glace étaient en partie reliées aux algues de glace. De plus, la thermodynamique de la couverture de glace peut contrôler la distribution des microorganismes et de la MO, comme observé lors de la fonte de la glace à ISPOL et lors du refroidissement de la banquise à SIMBA. La distribution des bactéries n’est pas corrélée avec celle des algues et de la MO dans la glace. En effet, la consommation de la MO par les bactéries semble être limitée non pas par la nature chimique des substrats mais par un facteur extérieur affectant le métabolisme bactérien tel que la température, la salinité ou une toxine. Le dysfonctionnement de la boucle microbienne menant à l’accumulation de la MO dans la glace a donc été mis en évidence dans nos échantillons.</p><p><p align="justify">De plus, le biofilm formé par les TEP est aussi impliquée dans l’attachement des cellules et autres composés aux parois des canaux de saumure et donc dans la séquence de largage lors de la fonte. Cette séquence semble propice au développement d’efflorescences phytoplanctoniques dans la zone marginale de glace. Les microorganismes originaires de la glace (surtout ceux de taille < 10 μm) semblent capables de croître dans la colonne d’eau et l’apport en nutriments organiques et inorganiques apparaît favorable à la croissance des microorganismes pélagiques.</p><p><p align="justify">Enfin, l’influence des activités hétérotrophes sur l’export de carbone et l’efficacité de la pompe biologique à carbone a été évaluée dans la couche de surface (0-100 m) et mésopélagique (100-700 m) de l’océan. Au contraire de la glace, les biomasses et activités bactériennes suivaient les distributions de la chlorophyll a et de la MO. Elles diminuent fortement en dessous de 100-200 m, néanmoins les valeurs intégrées sur la hauteur de la colonne d’eau indiquent que la reminéralisation de la MO par les bactéries dans la zone mésopélagique est loin d’être négligeable, spécialement dans une région dominée par les diatomées.</p> / Doctorat en Sciences agronomiques et ingénierie biologique / info:eu-repo/semantics/nonPublished

Thesis (PhD)-- Stellenbosch University, 2012. / ENGLISH ABSTRACT: Climate change and biological invasions are major threats to biodiversity. In particular, these threats are predicted to influence terrestrial systems in the sub-Antarctic, where significant ecosystem responses to both have already been seen. In this thesis, the sub-Antarctic Prince Edward Island group is used as a model system in which to investigate key questions relating to climate change and invasive species impacts. The island group comprises two islands, Marion (MI) and Prince Edward (PEI), both of which are experiencing rapid warming, yet have different invasive assemblages and in consequence are experiencing different impacts. Variation in the patterns of invasive species richness and abundance and their underlying causes are matters of considerable ecological and conservation significance. While an increase in thermal energy availability typically results in an increase in species richness, the mechanisms underlying these patterns are poorly understood. In Chapter 2 of this thesis, these relationships are explored for springtails, an important component of the soil fauna on Marion Island. Energy explains a large amount of the spatial variation in indigenous and invasive springtail species richness. Disturbance thresholds and stressful temperatures are more important than increased population sizes in determining this variation in species richness. As both indigenous and invasive springtail species richness and abundance are strongly related to temperature, a warming climate could have far-reaching consequences for these organisms. In particular, invasive species are predicted to be at an advantage relative to indigenous species under warming conditions. One species where this seems especially likely, given its physiological responses to experimental warming and drying, is the large invasive tomocerid, Pogonognathellus flavescens. Determining whether this will be the case depends on understanding the factors underlying its range limits and abundance structure. Moreover, few studies have sought to distinguish the causal basis of abundance structure and range limits, particularly for invasive species. Thus, in Chapter 3, local microclimate variables and physiological tolerances of the invasive springtail, P. flavescens (a habitat generalist), are examined. The results suggest that the species should be widely distributed across a range of habitats on MI. However, the springtail is restricted to indigenous Poa cookii tussock grassland habitats in the southeast. The current range limits are set by dispersal limitation (i.e. contingent absences) whilst abundance structure is a function of variation in soil substrate quality. However, over time, the widening distribution of P. cookii, as a consequence of a major management intervention (the eradication of feral cats), may enable P. flavescens to colonise all suitable areas. In Chapter 4, the focus changes to what has been considered the third major response to climate change, along with range and phenological responses - changing animal body sizes. Body size is one of the most significant and obvious features of animals and is of considerable ecological and physiological importance. A prediction of the temperature-size rule (TSR) is that with warming, body size of the weevil species on both MI and PEI should decline. However, predation by mice of the weevils on MI should fundamentally affect the pattern of such change, causing it to differ from neighbouring PEI, indicating synergistic impacts between climate change and invasions. Analysis of a 24-year data set indicates a decline in the body size of all weevil species on PEI with increasing temperature. However, on MI, a negative relationship between mean annual temperature and body size is found only for Palirhoeus eatoni, a species not eaten by mice. A possible explanation for the positive relationships found for the other species could be due to higher metabolic demands imposed on mice in colder years than in warmer ones. Any increase in predation coupled with a preference for larger sizes, which the mice clearly show, would lead to a decline in the mean size of the weevil species. Due to the relationship between body size and metabolic rate and the importance of the weevils in the islands’ food webs, changes to the body size of these organisms could have significant consequences for the island ecosystems’ functioning. The thermal environment experienced by organisms also has a direct effect on survival, growth and reproduction. The physiological response of organisms to rapidly changing climates is therefore a primary concern. Organisms may respond to variable environmental conditions through phenotypic plasticity as well as behaviour. Chapter 5 of this thesis shows that of the weevil species and populations investigated on MI, most display phenotypic plasticity, the form of which is in keeping with the ‘Hotter is Better’ hypothesis. This could be due to rare extreme temperature events and the advantage for the performance curves to incorporate high temperatures experienced in the environment. Mismatches between thermal optima and preferred temperatures displayed by all species could mean that these weevils are well equipped to cope with warming conditions on MI unless the prediction of an increase of rare extreme events such as extreme temperatures is realised. Rapidly changing climates and an increase in the introduction of non-indigenous species are issues of major conservation concern. This has increased the significance of studies on the impacts of these threats. However, this thesis shows that to understand such processes, it is essential that an integration of disciplines be undertaken. This thesis thus adopts a multidisciplinary approach and highlights key issues associated with both climate change and biological invasions. The patterns and predictions of species and community responses to these environmental changes are complex. Moreover, predicting such responses is likely to be problematic, especially as multiple factors will change concurrently and how these factors might change is unclear. This highlights the importance of long-term records for understanding organism responses to such changes. Furthermore, impacts on indigenous species are likely to be exacerbated by the predicted increase in the rate of introductions with climate change. This makes the case for preventing the dispersal of invasive species to new areas all the more important. / AFRIKAANSE OPSOMMING: Klimaatsverandering en indringer spesies is belangrike bedreigings vir biodiversiteit. In besonder word voorspel dat hierdie bedreigings terrestriële sisteme in die sub-Antarktiese sal beïnvloed, waar beduidende ekosisteem reaksie aan beide reeds gesien is. In hierdie tesis word die sub-Antarktiese Prince Edward eiland groep as a model sisteem gebruik om belangrike vrae met betrekking tot klimaatsverandering en die impak van indringer spesies te ondersoek. Die eiland groep bestaan uit twee eilande, Marion (ME) en Prince Edward (PEE), wat beide versnellende verwarming ervaar, maar tog verskillende indringer samestellings het en vervolgens verskillende impakte ervaar. Variasie in die patrone van indringerspesierykheid en vollopheid en hulle onderliggende oorsake is van aansienlike omgewings en bewarings betekenis. Terwyl 'n toename in die beskikbaarheid van energie tipies lei tot 'n toename in spesierykheid, word die onderliggend meganismes van hierdie patrone swak verstaan. In Hoofstuk 2 van hierdie tesis, word hierdie verhoudings vir springsterte ondersoek, 'n belangrike komponent van die grond fauna op ME. Energie verduidelik 'n groot hoeveelheid van die ruimtelike variasie in inheemse en indringende springstert spesierykheid. Versteuringsdrempels en stressvolle temperature is meer belangrik as die toename in bevolking groottes in die bepaling van hierdie variasie in spesierykheid. Aangesien beide inheemse en indringende springstert spesierykheid en vollopheid sterk verwant is aan temperatuur, kan 'n verwarmende klimaat verreikende gevolge vir hierdie organismes hê. In die besonder word voorspel dat indringerspesies bevoordeeld sal wees relatief tot inheemse spesies onder verwarmende toestande. Een spesie waar dit veral blyk om geneig te wees, gegewe sy fisiologiese reaksie tot eksperimentele verhitting en uitdroging, is die groot indringer tomocerid, Pogonognathellus flavescens. Om te bepaal of dit die geval sal wees, hang af van die begrip van die onderliggende faktore van sy voorkomsgrense en vollopheidstruktuur. Daarbenewens is daar min studies wat gepoog het om te onderskei tussen die veroorsakende basis van vollopheidstruktuur en voorkomsgrense, veral vir indringerspesies. Dus, in Hoofstuk 3, word plaaslike mikroklimaat veranderlikes en fisiologiese toleransies van die indringer springstert, P. flavescens ('n habitat generalis), ondersoek. Die resultate stel voor dat die spesie wyd verspreid moet wees oor 'n verskeidenheid van habitatte op ME. Maar, die springstert is beperk tot inheemse Poa cookii polle grasveldhabitatte in die suidooste. Die huidige voorkomsgrense word daar gestel deur verspreidingsbeperking (dws voorwaardelike afwesighede), terwyl vollopheidstruktuur 'n funksie is van die variasie in die grond substraat kwaliteit. Maar, die uitbreidende verspreiding van P. cookii as gevolg van 'n groot bestuursingryping (die uitwissing van wilde huiskatte), kan P. flavescens in staat stel om alle geskikte gebiede te koloniseer met verloop van tyd. In Hoofstuk 4 verander die fokus na wat as die derde groot reaksie op klimaatsverandering beskou word, saam met voorkoms en fenologiese reaksies - veranderende diere liggaamsgroottes. Liggaamsgrootte is een van die beduidendste en mees voor die hand liggende eienskappe van diere en is van aansienlike ekologiese en fisiologiese belang. 'n Voorspelling van die temperatuur-grootte-reël (TGR) is dat met verwarming, liggaamsgrootte van die snuitkewerspesies op beide ME en PEE sal afneem. Hoe ookal, predasie deur muise van die snuitkewers op ME moet fundamenteel die patroon van sodanige verandering op PEE beïnvloed, wat sinergistiese impakte tussen klimaatsverandering en indringings aandui. Die ontleding van 'n 24-jarige datastel dui aan op 'n afname in die liggaamsgrootte van alle snuitkewer spesies op PEI met ‘n toename in temperatuur. Maar, op ME is 'n negatiewe verhouding tussen die gemiddelde jaarlikse temperatuur en liggaamsgrootte net gevind vir Palirhoeus eatoni, 'n spesie wat nie deur die muise geëet word nie. 'n Moontlike verduideliking vir hierdie positiewe verhoudings wat gevind is vir die ander spesies kan wees as gevolg van hoër metaboliese eise op die muise in kouer jare as in warmer jare. Enige toename in predasie, tesame met 'n voorkeur vir groter mates, wat die muise duidelik wys, sou lei tot 'n afname in die gemiddelde grootte van die snuitkewer spesies. As gevolg van die verhouding tussen liggaamsgrootte en metaboliese tempo, sowel as die belangrikheid van die snuitkewers in die eilande se voedselwebbe, kan veranderinge in die liggaamsgrootte van hierdie organismes beduidende gevolge op die eiland ekosisteme se funksionering hê. Die termiese omgewing wat deur organismes ervaar word het ook 'n direkte invloed op oorlewing, groei en voortplanting. Die fisiologiese reaksie van organismes op vinnig veranderende klimate is dus 'n primêre bron van kommer. Organismes kan reageer op veranderlike omgewingstoestande deur fenotipiese plastisiteit sowel as gedrag. Hoofstuk 5 van hierdie tesis toon dat van die snuitkewerspesies en bevolkings wat ondersoek is op ME, die meeste fenotipiese plastisiteit vertoon, die vorm wat in ooreenstemming is met die ‘Warmer is Beter’ hipotese. Dit kan wees as gevolg van seldsame uiterste temperatuur gebeure en die voordeel vir die prestasie kurwes om hoë temperature wat ervaar word in die omgewing in te sluit. Mismatches tussen termiese optima en voorkeur temperature vertoon deur alle spesies kan beteken dat hierdie snuitkewers goed toegerus is om die verhitting op ME te hanteer, tensy die voorspelling van 'n toename van seldsame uiterste gebeure soos uiterste temperature gerealiseer word. Vinnig veranderende klimate en 'n toename in die bekendstelling van nie-inheemse spesies is kwessies van groot bewarings kommer. Dit het die betekenis van studies oor die impak van hierdie bedreigings verhoog. Hierdie tesis toon egter dat om sulke prosesse te verstaan, dit noodsaaklik is dat 'n integrasie van die dissiplines onderneem word. Hierdie tesis aanvaar dus 'n multi-dissiplinêre benadering en beklemtoon die belangrike kwessies wat verband hou met beide klimaatverandering en biologiese indringing. Die patrone en voorspellings van spesies en die gemeenskapsreaksies op hierdie omgewingsveranderinge is kompleks. Verder, die voorspelling van sodanige reaksies sal waarskynlik problematies wees, veral omdat verskeie faktore gelyktydig sal verander en hoe hierdie faktore kan verander is onduidelik. Dit beklemtoon die belangrikheid van lang termyn rekords vir die begrip van organisme reaksies op sulke veranderinge. Verder, die impak van inheemse spesies is geneig om te vererger deur die voorspelde toename in die tempo van bekendstellings met klimaatsverandering. Dit maak die taak vir die bestuur van die voorkoming dat indringerspesies nuwe gebiede bereik al hoe meer belangrik.

Der Antarktische Krill (Euphausia superba) ist eine Schlüsselart des Ökosystems des Südozeans und möglicherwese die biomassereichste Art der Erde. Klimawandelbedingte Lebensraumänderungen, die Expansion der kommerziellen Krillfischerei, sowie sich erholende Bartenwalpopulationen setzen Krillpopulationen jedoch zunehmend unter Druck. Daher besteht ein großer Bedarf, die Auswirkungen dieser verschiedener Stressoren auf Krillpopulationen zu verstehen und vorherzusagen. Dabei helfen Simulationsmodelle, die das Wachstum und die Entwicklung des Krills vorhersagen. Modelle sind vielseitige Instrumente, mit denen sich die Auswirkungen verschiedener Stressoren auf Krill in unterschiedlichen Szenarien bewerten lässt, was sie für Management- und Projektionsstudien besonders nützlich macht. In dieser Arbeit untersuchen wir Möglichkeiten die Robustheit von Modellen zu verbessern, die das Wachstum von Krillindividuen unter verschiedenen Umweltbedingungen vorhersagen. Zunächst haben wir ein bestehendes Wachstumsmodell erweitert, indem wir es um realistischere physiologische Prozesse ergänzt haben, was zu verbesserten Wachstumsvorhersagen führte. Anschließend haben wir in der Vergangenheit publizierte Wachstumsmodelle verglichen, um zu untersuchen, ob die Mechanismen des Krillwachstums verallgemeinerbar sind, da die Modelle für verschiedene Regionen entwickelt und mit unterschiedlichen Daten kalibriert wurden, regelmäßig aber außerhalb dieser Regionen verwendet werden. Diese Information ist wichtig für künftige Simulationsstudien, da eine regionale Spezifität die Modelle in ihrer großskaligen Anwendbarkeit limitieren würde. Die Wachstumsvorhersagen der verschiedenen Modellen bei identischen Inputdaten zeigte dabei, dass die Mechanismen und Wachstumsvorhersagen zwischen den Modellen erheblich variieren. Durch die Identifizierung der Ursachen dieser Modellspezifität, konnten wir das allgemeine Verständnis der Krill-Wachstumsmechanismen verbessern und zu der Entwicklung robusterer Modellen beitragen. Individuenbasierte Modelle ermöglichen die Einbindung vieler Prozesse, wie z. B. Verhalten, was diese Art von Modellen besonders attraktiv macht. Unser mechanistisches Verständnis des Verhaltens von Krill ist jedoch noch begrenzt, und wir haben versucht, dies durch die Analyse eines achtmonatigen Datensatzes über das vertikale Migrationsverhalten des Krills zu verbessern. Die Analysen zeigten, dass das vertikale Migrationsverhalten von Krillschwärmen sehr flexibel ist und wahrscheinlich von mehreren Faktoren wie dem Stadium, der Jahreszeit, den Nahrungsbedingungen und der Anwesenheit von Krillprädatoren bestimmt wird. Diese Studie dient als Grundlage für die Entwicklung von Modellkomponenten, die es erlauben das Krillverhalten in individualbasierte Modelle zu integrieren. Anschließend haben wir das vorhandene Wissen über das Krillverhalten in einem Reviewartikel zusammengefasst und dabei die möglichen Auswirkungen des Klimawandels auf Krillverteilung und das Schwarmverhalten analysiert. Zuletzt haben wir einen Systematic Literature Review durchgeführt, um eine Übersicht von Daten zu erstellen, die für die Modellentwicklung und -parametrisierung genutzt werden können. Auf diese Weise konnten wir Aspekte der Krillökologie identifizieren, für die aktuell besonders wenige Daten existieren, was wertvoll für zukünftige empirische Forschung sein wird. Da die Schließung dieser Datenlücken Zeit brauchen wird, diskutieren wir zudem auch Möglichkeiten zur Modellierung des Krillwachstums trotz bestehender Unsicherheiten. Mit dieser Arbeit leisten wir einen Beitrag zur Entwicklung realistischerer und robusterer individuenbasierter Krill-Wachstumsmodelle.:1. Introduction 2. Article I - The impact of seasonal regulation of metabolism on the life history of Antarctic krill (Euphausia superba) 3. Article II - An intercomparison of models predicting growth of Antarctic krill (Euphausia superba): the importance of recognizing model specificity 4. Article III - An open and lightweight method to analyze the vertical distribution of pelagic organisms using echogram screenshots 5. Article IV - Plasticity and seasonality of the vertical migratory behaviour of Antarctic krill using acoustic data from fishing vessels 6. Article V - Impacts of climate change on krill behaviour and population dynamics 7. Article VI - Mapping the data landscape for developing and parameterizing Antarctic krill models 8. Supervised Research Projects 9. Discussion 10. Literature cited in Introduction and Discussion 11. Appendices

This thesis describes medical aspects of the expeditions of the Heroic Age of Antarctic exploration (1895-1922). It is divided into three sections. Section 1 describes the drugs and medical equipment taken to the Antarctic by these expeditions. There is an introductory discussion followed by papers on ophthalmic drugs, injections, inhalations and suppositories, oral drugs and topical preparations. Sledging medical cases are considered separately. Brandy was often used as a medicine and the medical uses of alcohol are described. Some expeditions took what were described as “medical comforts”. This term was sometimes used as a euphemism for alcoholic beverages but, in fact, encompassed a wide range of foods for invalids. Burroughs Wellcome and Co. supplied many of the expeditions with drugs and their medical chests. They used the expedition link in their advertising and the relationship between the expeditions and drug companies is described. Section 2 describes some of the medical problems encountered. The most serious problems were scurvy and a condition often called “polar anaemia”, which seems to be the same as a condition known at the time as “ship beriberi” and what is now described as “wet beriberi”. The controversy as to whether Captain Scott and his colleagues died of scurvy is also discussed. Other problems included frostbite and snow blindness. At least 11 general anaesthetics were given, including two for amputation of frostbitten toes. Psychological problems were common and there was some serious psychiatric illness including alcohol abuse. Section 3 describes the doctors and some of the research that they carried out. The most common research done by doctors was bacteriological. Most doctors collected biological data on the explorers including weights and haemoglobin measurements. This was largely for health monitoring but one doctor pursued physiological research. Three doctors and a medical student studied geology.

Thesis (MSc)--Stellenbosch University, 2008. / ENGLISH ABSTRACT: Macroecological relationships have rarely been studied at wide spatial scales and across geographic ranges of species in the field in the sub-Antarctic. In this thesis I examined the occupancy, distribution and abundance, and the relationships thereof, of indigenous plants and alien species at broad (island-wide) and fine scales across sub-Antarctic Marion Island. The impacts of alien species and their interactions with indigenous plants were also investigated. I examined the nature of the abundance structure of a cushion-forming, vascular plant, Azorella selago, at the island-wide scale. Moreover, the hypothesis that species reach their highest abundances at the centre of their geographic range and decline in abundance towards the range edges was tested. Azorella selago cushions were counted in 8 m x 8 m quadrats, placed regularly at 1 minute latitude and longitude intervals across Marion Island. Using spatially non-explicit and explicit methods, this study showed that the abundance structure of A. selago had a more complex pattern of high abundance patches and low abundance gaps in its island-wide distribution. Subsequently, the hypothesis of an abundant centre distribution was not supported for A. selago across Marion Island. Rather, there were sharp discontinuities at both the coastal and altitudinal (667 m a.s.l.) limits for the species, between which little pattern in altitudinal abundance structure existed. Mice (Mus musculus) have recently been found to cause extensive structural damage to A. selago. The structural influence of mice on vegetation structure at the landscape scale has largely been overlooked on many sub-Antarctic islands. I mapped the distribution of evidence of mouse damage within the cushions of A. selago across the island using systematic (at 1 minute latitude and longitude intervals) and opportunistic sampling. Approximately 40 % of the systematically sampled sites had evidence of mouse damage to A. selago. Furthermore, a third of cushions in opportunistically sampled sites was damaged. Mouse damage was high in sites of low A. selago abundance, emphasizing that impacts of mice may be greater in low cushion abundance areas. This damage sometimes led to the disintegration of entire cushions. Given that A. selago acts as a nurse plant and supports high abundances of indigenous invertebrates, the impacts of mice on this keystone species may have significant ecological implications. Studies on interactions between alien and indigenous plants are limited within the sub-Antarctic. I examined the fine-scale distributions and co-occurrences of alien plants, Agrostis stolonifera and Sagina procumbens, and the indigenous Acaena magellanica (in 2 m x 2 m plots, subdivided into 0.25 m x 0.25 m quadrats) along rivers on Marion Island. Environmental variables were important for the occurrence of these species. In particular, 42.95 % and 24.82 % of the deviance in the occurrence of A. stolonifera and S. procumbens, respectively, was explained by environmental variables, compared to 17.35 % for A. magellanica. Furthermore, the co-occurrence of A. magellanica with A. stolonifera was significantly influenced by environmental variables. Significant positive spatial associations between A. magellanica and A. stolonifera were found, while the interactions of either species with S. procumbens were either spatially dissociated or random. Therefore, this study highlighted that alien species are responding to different environmental variables and conditions on Marion Island. Sagina procumbens seems to be less sensitive to the island’s environmental conditions and may thus be affecting biodiversity at broader ranges. This thesis provides unparalleled data on the distributions and interactions of indigenous plants and alien species for Marion Island. Alien species are undoubtedly posing significant threats to indigenous plants on the island and this thesis presents insight into interactions of species, specifically plants, an approach underrepresented in the sub-Antarctic to date. / AFRIKAANSE OPSOMMING: Makro-ekologiese verhoudings is selde bestudeer op wye ruimtelike skale en oor geografiese gebiede van spesies in die veld in die sub-Antarktiek. In hierdie tesis het ek die digtheid, bewoning, verspreiding en die verhoudings daarvan, van inheemse plante en uitheemse spesies ondersoek op breë (eiland-wyd) en smal skale oor sub-Antarktiese Marion Eiland. Die impakte van uitheemse spesies en hul interaksies met inheems plante is ook ondersoek. Ek het die aard van die digtheidstruktuur ondersoek van 'n kussing-vormende, vaatplant, Azorella selago, op die eiland-wydte skaal. Bowenal is die hipotese dat spesies hul hoogste digtheid bereik in die sentrum van hul geografies gebied en afneem in digtheid na die grense van die gebied getoets. Azorella selago kussings is getel in 8 m x 8 m kwadrante, wat eweredig geplaas is op 1 minuut breedte- en lengtegraad intervalle oor Marion Eiland. Deur gebruik te maak van ruimtelik nie-eksplisiete en eksplisiete metodes, het hierdie studie getoon dat die digtheidstruktuur van A. selago ‘n meer komplekse patroon van hoë digtheidslaslappe en lae digtheidsgapings in sy eiland-wyd verspreiding het. Vervolgens is die hipotese van 'n digte sentrum verspreiding nie gesteun vir A. selago oor Marion Eiland nie. Daar was eerder skerp diskontinuïteite by beide die kus- en hoërliggende (667 m bo seespieël) grense vir die spesie, waartussen daar ’n oneweredige patroon in digtheidstruktuur bestaan het. Onlangs is bevind dat muise (Mus musculus) uitgebreide struktuele skade aan A. selago veroorsaak. Die struktuele invloed van muise op plantegroei struktuur op landskapskaal is grootliks oor die hoof gesien op baie sub-Antarktiese eilande. Ek het die verspreiding van die bewyse van muisskade binne die kussings van A. selago oor die eiland gekarteer deur gebruik te maak van sistematiese (tot 1 minuut breedte- en lengtegraad intervalle) en opportunistiese opnames. Ongeveer 40 % van die sistematiese opname kwadrante het bewyse van muis skade aan A. selago vertoon. Verder, ‘n derde van die kussings in die opportunistiese opname kwadrante was beskadig. Muisskade was hoog in plotte met lae A. selago volopheid, wat beklemtoon dat impakte van muise groter mag wees in lae kussing digtheid gebiede. Hierdie skade het partykeer gelei tot die verbrokkeling van hele kussings. Gegewe dat A. selago as 'n verpleegsterplant optree en ’n hoë digtheid van inheemse invertebrata ondersteun, mag die impakte van muise op hierdie hoeksteen spesie beduidende ekologiese implikasies tot gevolg hê. Studies oor interaksies tussen uitheemse- en inheemse plante is beperk in die sub-Antarktiese gebied. Ek het die smal-skaal verspreidings ondersoek en medevoorkoms van uitheemse plante, Agrostis stolonifera en Sagina procumbens, en die inheems Acaena magellanica (in 2 m x 2 m kwadrante, subverdeel in 0. 25 m x 0. 25 m kwadrante) langs riviere op Marion Eiland. Omgewingsveranderlikes was belangrik vir die voorkoms van hierdie spesies. In besonder, 42.95 % en 24.82 % van die afwykings in die voorkoms van A. stolonifera en S. procumbens, onderskeidelik, is verduidelik deur omgewingsveranderlikes, vergeleke met 17.35 % vir A. magellanica. Verder, die medevoorkoms van A. magellanica saam met A. stolonifera is beduidend beinvloed deur omgewingsveranderlikes. Beduidende positiewe ruimtelike assosiasies tussen A. magellanica en A. stolonifera is gevind, terwyl die interaksies van beide spesies met S. procumbens was of ruimtelik nie-geassosieerd of lukraak. Daarom het hierdie studie uitgelig dat uitheemse spesies reageer op verskillende omgewingsveranderlikes op Marion Eiland. Sagina procumbens blyk minder sensitief te wees tot die eiland se omgewingsomstandighede en mag dus moontlik biodiversiteit op breër vlakke beïnvloed. Hierdie tesis voorsien onge-ewenaarde data oor die verspreiding en interaksies van inheems plante en uitheemse spesies vir Marion Eiland. Uitheemse spesies hou ongetwyfeld ’n beduidende bedreiging in vir inheems plante op die eiland, en hierdie tesis bied insig in die interaksies van spesies, spesifiek plante – ’n benadering wat swak verteenwoordig was in die sub-Antarktiese gebied tot op hede.

Thesis (MSc)--Stellenbosch University, 2011. / ENGLISH ABSTRACT: Species’ seasonal behaviour is of paramount importance in understanding community functioning and dynamics. Recently, plant phenology has further gained significance as a reliable indicator of climate change impacts. Despite the importance of understanding plant dynamics, there are relatively few plant phenological records for the sub-Antarctic region, and where records exist they are often not extensive. Sub-Antarctic Marion Island, typical of Southern Ocean Islands, offers a useful setting for addressing these knowledge gaps. This study documented the vegetative and reproductive phenologies (or aggregate phenological patterns) of twelve indigenous and three alien vascular plant species on the island. The phenological differences among the species and distinct seasonal groupings (e.g. early, intermediate and late species) were examined. I also investigated the phenological differences among the indigenous and alien plant species. Furthermore, the onset of selected reproductive phenophases from the current records was compared with historical records for determining the extent of climate change-related alterations in phenology. Phenological data were collected fortnightly on five, 5 m x 5 m permanent plots per species (except for a few species) for a full growing season. Thus the sample size is n = 5 for all plant species except for Crassula moschata (n = 4), Juncus effusus (n=4) and Rumex acetosella (n=1). Sites of the same species were separated by at least 500 m except for the alien plant, Juncus effusus, where all four known populations were selected despite two of these populations being < 500 m apart. This study indicated that Marion Island plants grow throughout the year with no major peaks except in Azorella selago and Acaena magellanica which showed winter dormancy. However, reproduction in most plant species predominately occurred in spring and summer months. Pringlea antiscorbutica and Poa cookii were the first two species to set flower buds in September while most species dispersed their seeds in summer except for Agrostis magellanica and Crassula moschata which dispersed in early autumn. Distinct from most temperate systems, the reproductive seasonality displayed by Marion Island plant species is explained more by daylength than by temperature, perhaps due to the region’s typical thermal aseasonality. Interestingly, many cooccurring species and/or clades across the Falkland, Kerguelen, Macquarie and South Georgia Islands also showed similar flowering onset date to the Marion Island plants, further confirming their daylength sensitivity. However, other external factors seem to come into play at later events of reproduction. Consequently, fruit maturation time of similar species across the sub-Antarctic islands varied substantially despite the plants having flowered in the same month. Although plant species showed similar reproductive seasonality, there were significant differences among species phenologies i.e. phenophase timing, duration and peak occurrence dates. However, using 95% confidence intervals of Generalized Linear Models weighted means, and/or one-way ANOVA (Tukey post hoc test), three homogenous sets of species (early, late, or intermediate onsets) were identified based on flower bud, flowering and seed dispersal phenophase onset dates. The homogenous species groupings observed for flower buds also remained unchanged during flowering onset except for Cotula plumosa and Callitriche antarctica which switched groups. As for the seed dispersal timing, the pattern was not consistent with that of the flower bud and flowering onset homogenous groupings, except for Acaena magellanica and Agrostis magellanica which remained in the early and late groups, respectively. Conversely, in the case of the timing of other phenophases (pollen release, fruit set and fruit ripening), entire phenophase durations, and peak occurrence dates, species overlapped greatly, resulting in an unbroken progression or continuum of phenology among species. Similarly, the three alien plant species investigated here (Cerastium fontanum, Juncus effusus and Rumex acetosella) showed no consistent phenological differences from the rest of the species. However, a widespread alien plant species on Marion Island, C. fontanum, reproduced for most of the year, although its reproduction peak was in summer months as was the case for the rest of the species. This study also indicated that indigenous plant species have altered their reproductive phenologies since 1965. Although the response was species-specific, the majority of plant species significantly delayed the onset of reproductive activities in 2007 by comparison with 1965. However, it is not clear if the observed species response was caused by the now drier and warmer Marion Island climate or by discrepancies in reporting in the earlier studies and/or sampling differences between the recent and historical records. Therefore, these results should be taken with caution. In conclusion, this research provided a detailed phenological dynamics record for vascular plant species on the island. Over time these records may be used as a basis for monitoring and modelling the impact of climate on plant phenology on the island. / AFRIKAANSE OPSOMMING: Spesies se seisoenale gedrag is van die allergrootste belang in die begrip van gemeenskapsfunksionering en dinamika. Meer onlangs het plant fenologie verdere betekenis verwerf as ‘n betroubare indikator vir die impakte van klimaatsverandering. Ondanks die belangrikheid om plant dinamika te verstaan, is daar relatief min plant fenologiese rekords vir die sub-Antarktiese streek en waar rekords wel bestaan is dit dikwels nie omvangryk nie. Sub- Antarktiese Marion Eiland, tipies van Suidelike Oseaan Eilande, bied ‘n nuttige ligging om hierdie kennis gapings aan te spreek. Hierdie studie het die vegetatiewe en voorplantingsfenologieë (of gesamentlike fenologiese patrone) van elf inheemse en drie uitheemse vaatplantspesies op die eiland gedokumenteer. Die fenologiese verskille tussen die spesies en duidelike seisoenale groeperings (bv. vroeë, intermediêre en laat spesies) is ondersoek. Ek het ook die betekenisvolle fenologiese verskille tussen die inheemse en uitheemse plantspesies ondersoek. Voorts, die aanvang van gekose voortplanting feno-fases van huidige rekords is vergelyk met historiese rekords om die mate van klimaatsverandering verbandhoudende veranderings in die fenologie te bepaal. Fenologiese data is twee weekliks ingesamel op vyf, 5 m x 5 m permanente plotte per spesie (behalwe vir ‘n paar spesies) vir ‘n volle groei seisoen. Dus, die insamelings grootte is n = 5 vir al die plantspesies behalwe vir C. moschata (n = 4), Juncus effusus (n=4) en Rumex acetosella (n=1). Persele vir dieselfde spesies is geskei deur ten minste 500 m, behalwe vir die uitheemse plant, Juncus effusus, waar al vier populasies wat bekend is gekies is, ten spyte daarvan dat twee van hierdie populasies < 500 m uitmekaar is. Hierdie studie het aangedui dat Marion Eiland plante regdeur die jaar groei, met geen belangrike spitstye nie, behalwe in Azorella selago en Acaena magellanica wat ‘n winter rusperiode wys. Hoe ookal, voortplanting in meeste van die plantspesies het hoofsaaklik voorgekom tussen die lente en somermaande. Pringlea antiscorbutica en Poa cookii was die eerste twee spesies om blomknoppe uit te stoot in September, terwyl die meeste spesies hulle sade versprei het gedurende die somer, behalwe vir Agrostis magellanica en Crassula moschata wat versprei het in vroeg herfs. Duidelik van meeste gematigde sisteme, word die voortplanting seisoenaliteit, getoon deur die Marion Eiland plantspesies, verduidelik meer deur daglengte as deur temperatuur, moontlik weens die streek se tipiese termiese a-seisoenaliteit. Interessant, baie spesies en/of afstameling-groeperings wat saam aangtref word dwarsoor die Falkland, Kerguelen, Macquarie en Suid Georgia Eilande wys ook soortgelyke bloei aanvangsdatums as die Marion Eiland plante, nog meer bevestigend van hulle dag-lengte sensitieweteit. Hoe ookal, ander eksterne faktore blyk betrokke te raak by latere gebeure van voortplanting. Gevolglik het vrug rypwordingstyd van dieselfde spesies oor die sub-Antarktiek noemenswaardig verskil, ten spyte daarvan dat die plante in dieselfde maand geblom het. Alhoewel plantspesies dieselfde voortplanting seisoenaliteit gewys het, was daar ‘n noemenswaardige veskil tussen spesie fenologieë, m. a. w. feno-fase tydsberekenning, tydsduur en spits voorkomsdatums. Hoe ookal, deur gebruik te maak van 95% betroubaarheid intervalle van Algemene Lineêre Modelle gewigte gemiddelde en/of een rigting ANOVA (Turkey post hoc toets), is drie homogene stelle van spesies (vroeë, laat en intermediêre aanvang) geïdentifiseer gebasseer op blomknop, bloei en saad verspreiding feno-fase aanvangsdatums. Die homogene spesie groeperings waargeneem op blomknoppe het ook onveranderd gebly gedurende bloei aanvang behalwe vir Cotula plumosa en Crassula antarctica wat groepe geruil het. Vir die saadverspreiding tydsberekenning was die patroon nie konstant met die van die blomknop en bloei aanvang homogene groepe nie, behalwe vir Acaena magellanica en Agrostis magellanica wat in die vroeë en laat groepe respektiewelik gebly het. Omgekeerd, in die geval van tydsberekenning van ander feno-fases (stuifmeel vrysetelling, vrugwerp, vrugrypwording), volledige feno-fase tydsduur en spits voorkomsdatums het spesies grootliks oorvleuel, wat ‘n ongebroke vordering of deurlopendheid van fenologie tussen die spesies tot gevolg het. Ooreenkomstig het die drie uitheemse spesies wat hier ondersoek is (Cerastium fontanum, Juncus effusus en Rumex acetosella) geen bestendige fenologiese verskille van die res van die spesies gewys nie. Hoe ookal, ‘n wydverspreide uitheemse spesie op Marion Eiland, Cerastium fontanum, het deur die meeste van die jaar voortgeplant, hoewel met ‘n voorplanting spits in die somer maande soos die res van die spesies. Hierdie studie dui ook aan dat inheemse plantspesies hulle voortplanting fenologieë verander het sedert 1965. Alhoewel die reaksie spesiespesifiek was, het die meerderheid van die plantspesies hulle voortplanting aanvang aansienlik vertraag gedurende 2007 in vergelyking met 1965. Hoe ookal, dis nie duidelik of die waargeneemde spesie reaksie was as gevolg van die nou droër en warmer Marion Eiland klimaat of deur teenstrydighede in verslagewing gedurende die vroëre studies en/of insameling verskille tussen die onlangse en historiese rekords. Daarom moet hierdie resultate met versigtigheid hanteer word. In samevatting, hierdie navorsing voorsien ‘n gedetaileerde fenologiese dinamieka rekord vir vaatplantspesies op die eiland. Oor tyd kan hierdie rekords gebruik word as basis vir monitering en modellering van die impak van klimaat.

In this thesis, I describe large-scale and small-scale features of the Antarctic Circumpolar Current (ACC) by merging conductivity-temperature-depth (CTD) data obtained by novel animal-borne sensors with data obtained by more conventional means. Twenty-one CTD-Satellite Relay Data Loggers (CTD-SRDLs) were attached to Southern elephant seals (Mirounga leonina) on South Georgia in 2004 and 2005. This was part of a larger international study (Southern Elephant Seals as Oceanographic Samplers; SEaOS), in which I played a major role in developing the oceanographic approach used to integrate physical data from a range of sources, and the means to link biological findings to oceanographic parameters. The development of animal-borne oceanographic sensors and their potential place within an ocean observing system is reviewed initially. Then, I describe the Series 9000 CTD-SRDL in detail, discussing its performance in the lab and during two field experiences with Southern elephant seals and Weddell seals (Leptonychotes weddellii ). Following this, a detailed study of the ACC frontal system in the South Atlantic is presented that uses merged Argo float data and CTD-SRDL data. The structure of the frontal field revealed by this unique dataset is examined, and unprecedented insight into its variability is obtained. Amongst the important findings is that, contrary to most climate models, our in situ data suggest a northward shift of the ACC east of 40W in 2004 and 2005 compared to previous work. Next, two CTD-SRDL sections are presented to identify the locations of the ACC fronts across Drake Passage, and an empirical relationship between upper ocean temperature and baroclinic mass transport is used to determine the transport through Drake Passage at the times of the sections. This technique is a powerful complement to more conventional means of data collection in this region, especially given the ability of the seals to conduct "sections" at times when ship-based fieldwork is logistically most challenging, i.e. in the winter time. The CTD-SRDLs do not only record hydrographic data, but simultaneously record seal movements and diving behavior. This enables insight to be obtained on the behavioral and physiological responses of Southern elephant seals to spatial environmental variability throughout their circumpolar range. The resulting energetic consequences of these variations could help explain recently observed spatially varying population trends. With a stable population at South Georgia and declining populations at Kerguelen and Macquarie Island. This study also highlights the benefits to the sensorcarrying animals themselves by showing the usefulness of this approach in examining the sensitivity of top predators to global and regional-scale climate variability. More importantly, I conclude that, by implementing animal-borne sensors into ocean observing strategies, we not only gain information about global ocean circulation and enhance our understanding of climate and the corresponding heat and salt transports, but at the same time we increase our knowledge about ocean’s top predators, their life history and their sensitivity to climate change.

1. Antarctic krill (Euphausia superba) form aggregations known as swarms that vary greatly in size and density. Six acoustic surveys were conducted as part of multidisciplinary studies at two study sites, the western and eastern core boxes (WCB and ECB), during the 1997, 1998 and 1999 austral summers, at South Georgia. A quantitative, automated, image processing algorithm was used to identify swarms, and calculate swarm descriptors, or metrics. In contrast to acoustic surveys of aggregations of other pelagic species, a strong correlation (r = 0.88, p = 0.02, 95% C.I.= 0.24 to 0.99) between the number of krill swarms and the mean areal krill density [rho.hat] was found. Multivariate analysis was used to partition swarms into three types, based on contrasting morphological and internal krill density parameters. Swarm types were distributed differently between inter-surveys and between on and off-shelf regions. This swarm type variation has implications for krill predators, by causing spatial heterogeneity in swarm detectability, suggesting that for optimal foraging to occur, predators must engage in some sort of adaptive foraging strategy. 2. Krill predator-prey interactions were found to occur at multiple spatial and temporal scales, in a nested, or hierarchical structure. At the largest inter-survey scale, an index of variability, I, was developed to compare variation in survey-scale predator sightings, sea temperature and [rho.hat]. Using I and a two-way ANOVA, core box, rather than year, was found to be a more important factor in determining species distribution. The absence of Blue-petrels (Halobaena caerulea) and the elevated number of Antarctic fur seals (Arctocephalus gazella) suggest that 1998 was a characterised by colder than average water surrounding South Georgia, and a high [rho.hat] in the ECB. At the smaller, intra-survey scales (<80 km, <5 day), the characteristic scale (distances in which predator group size, or krill density were similar, L_s) were determined. For krill and predators L_s varied by survey and the L_s of krill also varied by depth within a survey. Overlap in L_s were stronger between predator species than between a predator species and krill, indicating predators were taking foraging cues from the activity of predators, rather than from the underlying krill distribution. No relationship was found between swarm characteristics and predator activity, suggesting either there is no relationship between krill swarms and predators, or that the predator and acoustic observation techniques may not be appropriate to detect such a relationship. 3. To overcome the 2-D sampling limitations of conventional echosounders, a multibeam echosounder (MBE) observed entire swarms in three-dimensions. Swarms found in the nearshore environment of Livingston Island situated in the South Shetland Islands, exhibited only a narrow range of surface area to volume ratios or roughnesses (R = 3.3, CV = 0.23), suggesting that krill adopt a consistent group behaviour to maintain swarm shape. Generalized additive models (GAM) suggested that the presence of air-breathing predators influenced the shape of a krill swarm (R decreased in the presence of predators: the swarm became more spherical). A 2D distance sampling framework was used to estimate the abundance, N, and associated variance of krill swarms. This technique took into account angular and range detectability (half-normal, [sigma_r.hat] = 365.00 m, CV = 0.16) and determined the vertical distribution of krill swarms to be best approximated by a beta-distribution ([alpha.hat] = 2.62, [CV.hat] = 0.19; [beta.hat] = 2.41, [CV.hat] = 0.15), giving the abundance of swarms in survey region as [N.hat] = 5,062 ([CV.hat] = 0.35). This research represents a substantial contribution to developing estimation of pelagic biomass using MBEs. 4. When using a single- or split-beam missing pings occur when the transmit or receive cycles are interrupted, often by aeration of the water column, under the echosounder transducer during rough weather. A thin-plate regression spline based approach was used to model the missing krill data, with knots chosen using a branch and bound algorithm. This method performs well for acoustic observations of krill swarms where data are tightly clustered and change rapidly. For these data the technique outperformed the standard MGCV GAM, and the technique is applicable for estimating acoustically derived biomass from line transect surveys.

Outfacing the Storm is a song cycle that tells the story of the spectacular failure-turned-triumph of the Imperial Trans-Antarctic Expedition. Led by Sir Ernest Shackleton, the Expedition was an attempt to trek overland across the whole of the Antarctic continent. Shackleton’s ship, the Endurance, launched from England in August of 1914 with twenty-eight men. Within weeks of entering the polar latitudes, ice ensnared the ship. The Endurance remained stuck fast until the spring breakup crushed its hull, and it sank in November of 1915. The crew struck out for civilization, and, after six months and over one thousand miles of travel by foot, by ice-drift, and by lifeboat, they returned home alive. The Expedition’s safe return was heavily credited by its crew to Shackleton himself. A man of intense character, burning passion, and unfailing determination, Shackleton put the needs of his crew ahead of all else. His individual dedication to each member of the voyage was an inspiration to each. Ernest Shackleton, however, was a private man. He hid his own concerns from all except those closest to him. During the Expedition, he feared for the well-being of the crew. In the largest sense, he feared failure. Shackleton was only ever content while at sea. “Sometimes,” he wrote to his wife, “I think I am no good at anything but being away in the wilds… I grow restless and feel any part of youth is slipping away from me and that nothing matters… I feel I am no use to anyone unless I am outfacing the storm in wild lands.” This song cycle tells the tale of the Expedition through Shackleton’s eyes. The cycle is in eleven movements—nine recounting the story, plus a prologue and epilogue. All of the text is Shackleton’s own. The nine middle movements are taken from Shackleton’s memoir about the voyage, South. The prologue and epilogue are taken, respectively, from an interview and the above letter. / Music, Butler School of / text