Thesis (DPhil)--University of Stellenbosch, 2008. / ENGLISH ABSTRACT: Sub-Antarctic islands represent the only mid to high latitude terrestrial biomes in the Southern Hemisphere. These islands have various geological origins and histories, well-preserved terrestrial ecosystems and high levels of species endemism. In an attempt to understand the evolution and biogeography of terrestrial taxa in the South Polar Region, the first broad-scale molecular phylogeny was constructed for the unique terrestrial group, the ameronothroid mites (genus Halozetes (Oribatida)), collected from sub-Antarctic and Maritime Antarctic localities. Phylogenetic analyses based on a combined mitochondrial (cytochrome oxidase subunit I (COI)) and nuclear (histone-3 (H3)) sequence dataset indicated that the evolution of these mites were habitat specific (i.e. intertidal, supralittoral and terrestrial). Notwithstanding criticisms levelled against a molecular clock, the mites were evolutionary young (<10myo), contrary to their status as an ancient group predating Gondwana fragmentation. Biogeographic analyses indicated a complex pattern mainly sculpted by multiple independent dispersal events across the Antarctic Polar Frontal Zone similar to previous findings for other marine and terrestrial taxa. Also, the molecular phylogeny displayed considerable discourse with contemporary taxonomy suggesting the need for taxonomic revisions and reassessment of morphological characters. Sub-Antarctic Marion Island, the larger of the two islands comprising the Prince Edward Island archipelago (PEI), has experienced extensive glaciation and volcanism. To assess the impact of historical events (volcanism (including recent eruptions) and glaciation) and contemporary mechanisms (gene flow) on the genetic spatial distribution of species from Marion Island, two mite species namely Eupodes minutus (Prostigmata) and Halozetes fulvus (Oribatida) as well as a single plant species, Azorella selago (Apiaceae), were selected as model organisms. For independent phylogeographic analyses, mitochondrial sequence data (COI) were obtained for both mite species, while chloroplast sequence (trnH-psbA) and amplified fragment length polymorphism (AFLP) data were generated for the cushion plant, A. selago. Since A. selago is typified by two growth forms namely discrete cushions and continuous mats, it was essential to examine the growth dynamics prior to phylogeographic analyses. The sequence and fragment data indicated that both mite and plant species were significantly substructured across Marion Island. Manual comparisons indicated unique populations on the western (Kaalkoppie for H. fulvus, La Grange Kop for E. minutus and Mixed Pickle for A. selago), eastern (Bullard Beach for H. fulvus and Kildalkey Bay for E. minutus), northern (Middelman and Long Ridge for H. fulvus) and southern side (Grey Headed for H. fulvus and Watertunnel for A. selago) of the island. Importantly, the western side had unique localities for all species. Interestingly, based on the H. fulvus data, the western populations were relatively young, characterized by high migration rates, small effective (female) population sizes with no isolation-by-distance. The opposite scenario was found for the eastern populations. This spatial genetic structure described for species on Marion Island can be ascribed to both historical events and environmental conditions. These areas with their unique genetic composition are of special conservational concern; consequently this research will contribute to an active management plan for PEI, South Africa’s only Special Nature Reserve. / AFRIKAANSE OPSOMMING: Sub-Antarktiese eilande verteenwoordig die enigste terrestriële bioom in die middel tot hoër breedtegrades van die Suidelike Halfrond. Hierdie eilande besit ‘n verskeidenheid van geologiese oorspronge en geskiedenisse, goed-bewaarde terrestriële ekosisteme en hoë vlakke van endemisme. In ‘n poging om die evolusie en biogeografie van terrestriële taksa in die Suid Pool Area te verstaan, is die eerste grootskaalse molekulêre filogenie saamgestel vir ‘n unieke terrestriële groep, die ameronothoïed miete (genus Halozetes (Oribatida: Ameronothroidea)), vanaf menigte sub-Antarktiese en Maritime Antarktiese lokaliteite. Filogenetiese analises gebaseer op die saamgestelde mitochondriale (sitokroom oksidase subeenheid I (COI)) en nukluêre (histoon-3 (H3)) basispaarvolgordes het aangedui dat die evolusie van hierdie miete habitat spesifiek is (m.a.w inter-gety, supralitoraal en terrestrieël). Ongeag die kritiek teenoor ‘n molekulêre klok, is hierdie miete evolusionêr jonk (<10mjo), wat teenstrydig is met hulle status as ‘n antieke groep wat terugdateer voor Gondwana fragmentasie. Biogeografiese analises het ‘n komplekse patroon aangedui wat grotendeels gekarakteriseer word deur menigte onafhanklike verspreidingsgebeurtenisse bo-oor die Antarktiese Polêre Frontale Zone, wat ooreenstemmend is met vorige bevindinge vir ander mariene en terrestriële taksa. Die molekulêre filogenie het ook aansienlik verskil van die tradisionele taksonomie, dus is taksonomiese aanpassings en herklassifisering van morfologiese karakters noodsaaklik. Sub-Antarktiese Marion Eiland, die groter eiland van die Prins Edward eilandgroep (PEI), het uitermate glasiasie en vulkanisme ondervind. Om die impak van historiese gebeurtenisse (vulkanisme (insluitend onlangse uitbarstings) en glasiasie) en kontemporêre meganismes (geenvloei) op die genetiesgespasieërde verspreiding van spesies vanaf Marion Eiland te bepaal, was twee mietspesies naamlik Eupodes minutus (Prostigmata) en Halozetes fulvus (Oribatida) asook ‘n enkele plantspesie, Azorella selago (Apiaceae), gekies as model organismes. Vir onafhanklike filogeografiese analises, was die mitochondriale basispaarvolgorde (COI) vir beide mietspesies bepaal, terwyl chloroplast basispaarvolgorde (trnH-psbA) asook geamplifiseerde fragmentlengte polimorfisme (AFLP) data gegenereer was vir die kussingplant, A. selago. Aangesien A. selago gekenmerk word deur twee groeivorme, naamlik diskrete kussings en aaneenlopende matte, was dit noodsaaklik om eers die groeidinamika van die plant te ondersoek alvorens ‘n filogeografiese studie kon geskied. Die basispaarvolgordebepalings en fragmentdata het aangedui dat beide mietspesies sowel as die plantspesie betekenisvolle substruktuur vertoon regoor Marion Eiland. Informele vergelykings het unieke populasies aangedui op die westelike (Kaalkoppie vir H. fulvus, La Grange Kop vir E. minutus en Mixed Pickle vir A. selago), oostelike (Bullardstrand vir H. fulvus en Kildalkeybaai vir E. minutus), noordelike (Middelman en Long Ridge vir H. fulvus) en suidelike kant (Grey Headed vir H. fulvus en Watertunnel vir A. selago) van die eiland. Die westelike kant besit dus unieke lokaliteite vir al die spesies. Interressantheidhalwe het die H. fulvus data getoon dat die westelike populasies relatief jonk is en gekarakteriseer word deur hoë migrasiesyfers en klein effektiewe (vroulike) populasiegroottes met geen isolasie-oor-afstand nie. Die resultate vir die populasies aan die oostelike kant van die Marion Eiland was presies teenoorgesteld. Dié beskryfde substruktuur vir die spesies op Marion Eiland is afkomstig van beide historiese gebeurtenisse asook omgewingstoestande. Hierdie areas met hul unieke genetiese samestelling, is belangrik vir natuurbewaring. Hierdie navorsing sal bydra tot die bestuursriglyne van PEI, Suid Afrika se enigste Spesiale Natuurreservaat.

Thesis (PhD (Conservation Ecology and Entomology))--Stellenbosch University, 2008. / ENGLISH ABSTRACT: Shifts in species ranges are an important consequence of climate change, and can affect the composition, structure and functioning of ecosystems. Generally, in response to rising temperatures, species have increased their altitudinal and latitudinal distributions along their cooler boundaries, although there has been large variation between species. However, in addition to climatic factors, species range limits are also sensitive to biotic interactions. Therefore, interspecific interactions may be able to influence the impact of climate change on species distributions. In this thesis climate change, range shifts and spatial variation in plant-plant interactions are documented to examine the potential for biotic interactions to mediate climate-driven altitudinal range shifts on sub-Antarctic Marion Island. The climate on Marion Island changed considerably between 1949 and 2003, with significant trends in biologically-relevant parameters encompassing average climatic conditions, variability in climate parameters and extreme weather events. Combining these trends showed that while environmental conditions have ameliorated for some species, they have become more abiotically stressful for others (e.g. hygrophilous species). Concurrent with changes in the island’s climate, there have been rapid changes in the elevational distribution of Marion Island’s native vascular flora. On average species ranges expanded upslope (as predicted in response to the warming experienced on the island), although individual range expansion rates varied greatly. As a result of the idiosyncratic expansion rates, altitudinal patterns of species richness and community composition changed considerably. Therefore, both speciesand community-level changes occurred in the flora of Marion Island, demonstrating the community’s sensitivity to climate change. To determine the potential for interspecific interactions to have mediated these changes in species elevational ranges, spatial variation in the balance of positive and negative plant-plant interactions were examined. Increasing spatial association between four dominant species along an altitudinal severity gradient suggested that the intensity of facilitation, relative to the intensity of competition, is greater under more severe conditions. While, interaction strength varied between species pairs, the nature of relationship was consistent across the species. At a broader spatial scale, the performance of the grass Agrostis magellanica was compared in the presence and in the absence of the cushion plant Azorella selago along the entire altitudinal range of both species. The influence of the cushion plant on A. magellanica switched from negative to positive with increasing environmental severity, reaching a positive asymptote under extreme environmental conditions. Therefore, on Marion Island the spatial variation in the nature of biotic interactions is related to environmental severity, and facilitative interactions can be strong under extreme environmental conditions. These results show that positive biotic interactions are important at higher elevations on Marion Island, and could thus accelerate upslope range expansions in response to climate change. Nonetheless, competitive inhibition of upslope species movement, especially in areas of dense vegetation (i.e. mid or low altitudes), could have the opposite result, acting antagonistically with the effect of rising temperatures. This research highlights the importance of considering interspecific interactions when examining the biotic implications of climate change, both for affecting the rate at which species ranges change and the extent of species distributions. / AFRIKAANSE OPSOMMING: Die verskuiwing van spesies se verspreidingspatrone is ‘n belangrike gevolg van klimaatsverandering en kan die samestelling, struktuur en funksionering van ekosisteme affekteer. Oor die algemeen, in reaksie op verwarming, kom spesies op hoër hoogtes bo seespieël en hoër breedtegrade voor, alhoewel daar groot verskille tussen spesies in hierdie aspek is. Behalwe vir klimatologiese faktore, is spesies se verspreidingslimiete ook sensitief vir biotiese interaksies met ander spesies. Daarom kan interaksies tussen spesies die effek van klimaatsverandering op spesiesverspreiding verder beïnvloed. In hierdie verhandeling word klimaatsverandering, verandering in verspreiding van spesies en geografiese variasie in plant-plant interaksies ondersoek. Dit word gedoen ten einde die moontlikheid te toets dat biotiese interaksies die verandering in verspreidingspatrone van klimaat gedrewe spesies op sub-Antarktiese Marion Eiland beïnvloed. Marion Eiland se klimaat het aansienlik tussen 1949 en 2003 verander, met betekenisvolle neigings in biologies-relevante parameters, insluitend gemiddelde klimaatstoestande, variasie in klimaat en uiterste weerstoestande. As die veranderinge saam beskou word, wys dit dat, terwyl omgewingstoestande vir sommige spesies verbeter het, toestande vir ander spesies abioties meer ongunstig geword het (bv. akwatiese plantsoorte). Tesame met die veranderinge in die eiland se klimaat, het die hoogte bo seespieël verspreiding van Marion Eiland se inheemse vaatplante vinnig verander. Gemiddeld het spesies-verspreiding se hoogte bo seespieël na hoër hoogtes verskuif (soos voorspel in reaksie op verwarming), alhoewel die tempo van verskuiwing na hoër hoogtes tussen spesies verskil het. As gevolg van hierdie idiosinkratiese reaksies, het die aantal en samestelling van plant spesies op verskillende hoogtes op die eiland aansienlik verander. Die sensitiwiteit van Marion Eiland se flora ten opsigte van klimaatsverandering word deur hierdie verandering gedemonstreer. Om te bepaal of die interaksies tussen spesies verspreidingspatrone kan beïnvloed, is die geografiese variasie in die balans van positiewe en negatiewe interaksies ondersoek. Vier dominante vaatplante het meer geassosieerd geraak hoe hoër hulle bo seespieël voorgekom het. Dit is ‘n aanduiding dat die sterkte van die positiewe interaksies (fasilitering), relatief tot die sterkte van die negatiewe reaksie (kompetisie), groter onder uiterste omgewingstoestande is. Die sterkte van die interaksie het verskil tussen spesies pare, maar die verhouding tussen omgewingstoestande en die sterkte van die interaksie was dieselfde vir alle spesies. Op ‘n breër geografiese vlak is die opbrengs van die gras Agrostis magellanica in die teenwoordigheid en afwesigheid van die polsterplant Azorella selago op alle hoogtes waar die plante saam voorkom vergelyk. Die invloed van A. selago op A. magellanica was negatief op lae hoogtes bo seespieël, maar het meer positief geraak met ‘n toename in hoogte bo seespieël, tesame met omgewingshardheid. Die invloed van die polsterplant op die gras het ‘n positiewe asimptoot onder uiterste omgewingstoestande bereik. Dus, op Marion Eiland is die geografiese variasie in biotiese interaksies verwant aan omgewingstoestande, en positiewe interaksies kan selfs onder die uiterste omgewingstoestande sterk wees. Hierdie resultate wys dat positiewe biotiese interaksies tussen plante belangrik is by hoër hoogtes bo seespieël op Marion Eiland. Die interaksies kan dus die opwaartse verspreiding van spesies in reaksie op klimaatsverandering versnel. Nogtans kan negatiewe interaksies die teenoorgestelde effek hê aangesien kompetisie tussen plante, veral in plekke met digte plantegroei (d.w.s. lae of middel hoogtes bo seespieël) opwaartse verskuiwing van spesies kan verhinder. Hierdie navorsing dui aan hoe belangrik dit is om interaksies tussen spesies in ag te neem, aangesien die interaksies die tempo en omvang van veranderinge in verspreiding kan beïnvloed.

Thesis (PhD)--Stellenbosch University, 2012. / ENGLISH ABSTRACT: This study addresses a little-known but important part of South Africa’s history: its involvement with the sub-Antarctic and Antarctic in the twentieth century. It has a three-fold approach. Firstly, it provides insight into the motives driving South Africa’s investment in the region, from the first call for a South African Antarctic expedition in 1919 to the post-apartheid recommitment to the South African Antarctic Programme. Interrogating of the reasons behind South Africa’s activities in this region – including those that failed –throws into relief broader issues about how and where South Africa saw itself in the geopolitical order. As such, this dissertation is situated within a body of Antarctic scholarship that seeks to subvert the prevailing homogenising narrative of the continent as simply the preserve of scientists and heroes. In particular, it investigates how tropes of imperialism and nationalism functioned in these remote corners of the world. Secondly, this dissertation investigates how changing perceptions of the extreme environment of Antarctica, and specifically the Prince Edward Islands, can add to our understanding of environmental history. It also shows how the values projected onto and invested in the environment as ‘nature’ changed over time. Thirdly, it takes into account the humans that were South Africa’s presence in the region and how the underlying patterns in the fabric of South African society, including race and gender, crystalized on the Antarctic continent. / AFRIKAANSE OPSOMMING: Hierdie studie is gerig op ʼn minder bekende, maar belangrike aspek van Suid-Afrika se geskiedenis: die land se betrokkenheid by die sub-Antarktiese gebied en Antarktika in die twintigste eeu. Die studie volg van ʼn drie-ledige benadering. Eerstens, verskaf dit insig in die dryfvere agter Suid-Afrika se investering in die streek – vanaf die eerste beroep op ʼn Suid-Afrikaanse Antarktiese ekspedisie in 1919, tot die post-apartheid regering se herverbintenis tot die Suid-Afrikaanse Antarktiese program. Die ondersoek na die redes vir Suid-Afrika se aktiwiteite in die streek – insluitend dié wat misluk het – bring breër kwessies oor Suid-Afrika se selfbeskouing in die wêreld se geopolitieke orde, na vore. Hierdie studie word binne ʼn kritiese raamwerk van navorsing oor Antarktika geplaas. Dié raamwerk streef daarna om die oorheersende homogene beeld van die kontinent as die eksklusiewe grondgebied van wetenskaplikes en helde, onder die soeklig te stel. In die besonder stel dit ondersoek in na hoe imperialisme en nasionalisme in hierdie verafgeleë uithoeke van die aarde versinnebeeld is. Tweedens, ondersoek hierdie studie hoe veranderende persepsies van Antarktika - en veral die Prins Edward eilande - se uiterste omgewing tot ons begrip van omgewingsgeskiedenis kan bydra. Dit dui ook aan die mate waartoe bestaande waardes wat op die omgewing as ‘natuur’ geprojekteer en gevestig is, mettertyd verander het. Die derde benadering neem die mense wat Suid-Afrika se teenwoordigheid verpersoonlik het in aanmerking - en hoe die onderliggende patrone in die Suid-Afrikaanse samelewing, insluitend ras en geslag, op die Antarktiese kontinent uitgekristalliseer het.

All previous taxonomic studies on the Ectemnorhinus group of weevils have been based primarily on morphological data. While these studies are invaluable, some questions can only be addressed adequately through molecular studies. This is especially true when studying the genetic relationships and phylogeograpic patterns of taxa endemic to the South Indian Ocean Province (SIP) biotas that have long been controversial. The Ectemnorhinus group of genera is a monophyletic unit of weevils endemic to the region. The present study focused mainly on the Ectemnorhinus group of weevils found on the Prince Edward Islands archipelago (PEIA). The mitochondrial cytochrome oxidase I gene was targeted when investigating relationships among members of this weevil group. On the PEIA, it is important to note that Marion Island (MI) and Prince Edward Island (PEI) differ in terms of alien invasive species, such as the introduced house mouse Mus musculus and in conservation management strategies. Since emergence, a series of volcanic and glaciation events have occurred on Marion Island, whilst Prince Edward Island has remained largely unaffected by glaciation. Phylogenetic analyses revealed the presence of two genetically and morphometrically distinct species of Ectemnorhinus weevils on PEI, whilst evidence for a single species, comprising diverse genetically discrete populations was found on MI. Based on these results, the species unique to PEI has been designated E. kuscheli n. sp., whilst the present study confirmed the synonymy between E. similis and E. marioni, the two species originally described from MI. Ectemnorhinus kucheli appears to be restricted to PEI, whereas E. similis occurs on both MI and PEI. When investigating the population dynamics of the Ectemnorhinus weevils on the PEIA, the data indicated that PEI was the first of the two islands of the PEIA to be colonized by Ectemnorhinus weevils, at an estimated time of coalescence of approximately 0.3116 million years ago (MYA). The PEI population then acted as the source population for the colonization of MI by Ectemnorhinus weevils some time before the last glaciation, approximately 10 000 to 35 000 years ago. The separation by distance of the PEI Ectemnorhinus weevils from those on MI then gave rise to two species by allopatric speciation on MI. During the last glaciations, MI was extensively glaciated with only the southwestern corner of the island being free of ice. This extensive glaciation of MI would have resulted in the eradication of all E. similis on MI except for those occurring on the ice-free southwestern corner of the island. At the end of the last glacial maximum, when the ice started to melt, the coastal areas of MI emerged first from beneath the ice and were available for re-colonization by weevils. The movement of weevils that were isolated in the south-western corner of MI, along the coastal areas of the island, was assisted by strong, frequent south-western winds. Subsequent, post-glacial volcanism during the Holocene was then responsible for the fragmentation of the new migrants, resulting in small population pockets surrounded by fresh, uninhabitable lava and subsequent divergence of each populations. When the Holocene black lava became re-colonizeable, the weevils from the different isolated populations migrated to the remainder of the island. Currently, members of the different genetically-identified populations occur in sympatry and in some cases even on the same plant, but no noticeable geneflow was detected between them. It is thus suggested that the time of isolation, before the post-glacial black lava during Holocene became hospitable, was sufficiently long and the populations sufficiently small that a number of genetically-discrete populations arose. Consequently, the present study recognises two genetically discrete populations of E. kucheli on PEI and seven discrete E. similis populations on MI that are morphologically indistinct. When examining the relationships among 13 species from five different islands within the South Indian Ocean Province (SIP) that are representative of 22 populations within the genera Palirhoeus, Bothrometopus and Ectemnorhinus, there was little support for separating the genus Palirhoeus from Bothrometopus, and no support for the morphologically-delineated species groups currently recognized within Bothrometopus. The present study shows that colonization of the Prince Edward Islands is likely to have occurred repeatedly from other islands within the SIP and that Bothrometopus parvulus on the PEIA comprises two species that are not sister taxa. The second novel con-generic species was therefore designated Bothrometopus huntleyi n. sp. and examination of the genetically identified specimens resulted in the indentification of distinguishing morphological characteristics. The analyses indicated that B. huntleyi arose approximately 0.5 million years ago from a high-altitude population that is still present on MI. The first major intra- and inter-island dispersal event occurred ~0.338 MYA, coinciding with the glaciation-free second volcanic stage on MI. Apart from this early inter-island colonisation, only one other between-island dispersal event, corresponding with the glaciation-free seventh volcanic stage, was detected. Genetically discrete weevil complexes on each of the islands of the PEIA together with the low levels of inter-island gene flow reaffirm the need to control alien invasive mice, which are restricted to MI, and which prey on these weevil species. / Thesis (PhD)--University of Pretoria, 2012. / Zoology and Entomology / unrestricted

Killer whales are the oceans’ apex predator and are known to have important effects on ecosystems. At Subantarctic Marion Island, southern Indian Ocean, they have only been studied opportunistically, resulting in limited knowledge of their ecosystem impact here. This dissertation describes the prey and seasonal abundance, estimates the population size and assesses the predatory impact of killer whales on seals and penguins at Marion Island, using dedicated and opportunistic shore-based observations and photographic identification, from 2006 to 2009. During 823 sightings of killer whales at Marion Island (2006 to 2009) 48 predation events were recorded; in only 10 cases could prey be identified. Killer whales fed on fur seals, elephant seals and penguins. Constant effort (dedicated) observations (259 hours, 2008 to 2009) showed that killer whale abundance, which peaked in September to December with a secondary peak in April to May, is linked to the abundance of seals and penguins. Mark-recapture analyses were performed using nearly 10 000 photographs taken from 2006 to 2009. Following careful quality control criteria 37 individuals were identified and a population size of 42 (95% CI = 35-50) individuals estimated using the open population POPAN parameterization in the software program MARK. The analytical approach is more rigorous than that used in any previous population size assessment at Marion Island. Finally, the above data were integrated to assess whether top-down control of seal and penguin populations at Marion Island is generally plausible using a simple process of elimination. Based on published data I predicted the energetic ingestion requirements of adult male and female killer whales as 1 394 MJ.day-1 and 1 028 MJ.day-1, respectively. Expanding these requirements to the 37 killer whales photographically identified at Marion Island, the population requires 40 600MJ.day-1. Based on available energy density and mass data, I predicted the energy content of available seal and penguin prey and calculated the rates at which killer whales would consume these prey in various scenarios. Penguins and Subantarctic fur seals are relatively insensitive to killer whale predation owing to their large population sizes (10 000s to 100 000s), conversely, the smaller populations (100s to 1 000s) of Antarctic fur seals and southern elephant seals are sensitive to predation, particularly the latter as they have a high energy content (approximately 2 000 to 9 000 MJ). Populations of these seals are currently increasing or stable and I conclude that presently killer whale predation is not driving population declines, although they clearly have the potential for regulation of these smaller populations. Thus, if population sizes were reduced by bottom-up processes, if killer whale diet shifted, or if prey availability changed, top-down control by killer whales could become significant. This study provides baseline information for the informed management and conservation of killer whales at Marion Island, identifies avenues for further research, and provides a foundation for the continuation of structured and dedicated killer whale research at Marion Island. / Dissertation (MSc)--University of Pretoria, 2011. / Zoology and Entomology / unrestricted

Please read the abstract in the section 00front of this document. / Dissertation (MSc)--University of Pretoria, 2010. / Zoology and Entomology / unrestricted