MSc., Faculty of Science, University of the Witwatersrand, 2011 / This research investigated the susceptibility of South African indigenous aquatic plants to competition from invasive species, using the competitive interactions of two sets of aquatic plants as a potential indicator. These interactions were studied in two separate experiments: the submersed weeds, Hydrilla verticillata (L.F.) Royle (Hydrocharitaceae) and an indigenous species, Lagarosiphon major (Ridley) Moss (Hydrocharitaceae) and; the floating weeds, Azolla filiculoides Lamarck (Azollaceae) and the indigenous Spirodela polyrhiza (L.) Schleid (Lemnaceae). Plants were grown under differing nutrient levels, and in an addition series of eight different densities, using the reciprocal yield model to estimate competitive ability. The invasive Hydrilla outcompeted Lagarosiphon in terms of mean length, dry mass, and survival. Major algal infestation in the high nutrient level of the Hydrilla/Lagarosiphon experiment altered light and nutrient conditions, which may have played a significant role in the lack of establishment of Lagarosiphon and the poor growth performance of Hydrilla. The invasive Azolla and indigenous Spirodela both performed well in terms of plant mass and increase in number. While Azolla was affected by intraspecific competition, it showed a steady increase in growth and multiplication with an increase in nutrients. The individual mass of Spirodela plants was highest in the low nutrient level, and multiplication rates were greatest in the high nutrient level. Results indicate that the susceptibility of indigenous plants may be increased in highnutrient systems, and that a continuous monitoring programme of aquatic alien species is vital in protecting our indigenous plants from extinction. This research recommends that the method of investigating competitive interactions between alien and indigenous plants be repeated with a variety of aquatic plants, as a means of anticipating susceptibility to invasions.

Thesis (Ph. D.)--University of Canterbury, 2008. / Typescript (photocopy). Includes bibliographical references (leaves 148-172). Also available via the World Wide Web.

Thesis (Ph. D.)--Ohio State University, 2009. / Title from first page of PDF file. Includes bibliographical references (p. 151-168).

Thesis (M.Com. (Economics & Economic History)) - Rhodes University, 2008.

Thesis (M.S.)--Michigan State University. Dept. of Fisheries and Wildlife, 2006. / Title from PDF t.p. (viewed on June 19, 2009) Includes bibliographical references. Also issued in print.

Thesis (Ph. D.)--Georgia Institute of Technology, 2006. / Title from PDF t.p. (viewed on Apr. 12, 2007). Streelman, J. Todd, Committee Member ; Miller, Margaret, Committee Member ; Weissburg, Marc, Committee Member ; Snell, Terry, Committee Member ; Hay, Mark, Committee Chair. Includes bibliographical references (p. 104-113).

La variabilité au sein des populations naturelles est un facteur primordial de maintien de la diversité fonctionnelle dans les communautés et les écosystèmes. Présente dans une aire géographique restreinte, la truite marbrée (Salmo marmoratus) est une espèce aujourd'hui menacée par les effets du changement climatique (augmentation de la fréquence et de l'intensité des crues) et par les invasions biologiques. Seules quelques populations persistent dans le bassin adriatique. Les objectifs de cette thèse sont de quantifier la variabilité trophique de cette espèce dans des populations naturelles de l'échelle populationnelle à l'échelle individuelle et d'en identifier les mécanismes. Le régime alimentaire de 1523 individus issus des sept populations du bassin de la Soca (Slovénie) échantillonnées entre 2011 et 2013 a été analysé à l'aide des isotopes stables (d15N et d13C). Les résultats mettent en évidence une forte variabilité trophique entre les individus et au sein des individus au cours de leur ontogénie. Cette variabilité trophique également observée à l'échelle populationnelle, dépend d'une part de la structure en taille des populations et d'autre part des effets des variables environnementales à la fois sur la physiologie des organismes et sur la disponibilité des ressources alimentaires. Les résultats indiquent un fort taux de cannibalisme chez cette espèce (de 6 à 32 % à l'échelle de la population) survenant par opportunisme et favorisé par la densité des proies. Le cannibalisme est un comportement permanent qui va affecter la dynamique des populations. L'étude des interactions trophiques entre la truite marbrée et une truite exotique (la truite arc-en-ciel, Oncorhynchus mykiss) montre que la compétition entre ces deux Salmonidés engendre un déplacement de niches trophiques de l'espèce native. Cependant, nos résultats suggèrent que le comportement piscivore de la truite marbrée, rarement observé chez des Salmonidés de rivière, confère à l'espèce une résistance biologique face aux invasions biologiques. L'ensemble des résultats démontre la forte plasticité trophique, à la fois individuelle et populationnelle, de ce prédateur aquatique, qui pourrait constituer un avantage adaptatif dans un contexte de changement climatique. / Variability within natural populations plays a major role in maintaining functional diversity in communities and ecosystems. The marble trout (Salmo marmoratus) is a species of great conservation concern given its restricted geographical area. Few populations persist in the Adriatic basin, threatened by the impacts of global change (increase of the frequency and intensity of flood events) and by biological invasions. In this thesis, we aimed to quantify trophic variability of this species both at the population and individual levels and to understand the underlying mechanisms. The diet of 1523 marble trout from the seven remnant populations persisting in the Soca River basin (Slovenia) were sampled between 2011 and 2013 and were analyzed with stable isotopes (d15N and d13C). Our results show strong variability through ontogeny within as well as between individuals. This intraspecific variation was observed at the population level and was determined by both environmental variables and size-structured demography effects on fish physiology and ecological opportunities. Results indicate a high rate of opportunistic cannibalism (from 6 to 32 % in populations) promoted by prey densities. This permanent behavior may lead to modification of the populations' size structures and dynamics. Studying trophic interactions between the marble trout and an invasive salmonid species (rainbow trout, Oncorhynchus mykiss), we showed that trophic niches of the native trout shifted because of interspecific competition. However, our results suggest that the piscivorous behavior of the marble trout, rare for stream-dwelling salmonids, would be a strong advantage for biotic resistance against biological invaders. This thesis demonstrates the high trophic plasticity, both at the individual and population levels, in this freshwater predator that could be an adaptative advantage in a global change context.

L’empreinte humaine sur Terre est si profonde qu’elle entraine des changements environnementaux qui affectent et modifient le fonctionnement des écosystèmes. Parmi tous les biotopes, les écosystèmes aquatiques continentaux sont des habitats remarquables qui abritent une grande biodiversité, mais qui sont aussi les plus menacés par les activitéshumaines. Les principales causes de la perte de qualité de l'eau et de la perturbation des ces écosystèmes comprennent la pollution des eaux et l'introduction d'espèces exotiques. La partie première partie de ce travail de thèse a montré des invasions parasitaires sur les populations de tortues d'eau douce indigènes, transmis par des espèces de tortues introduites. De plus, la nécessité d'étudier et de décrire les nouvelles espèces de parasites invasives avec des techniques plus performantes a été soulignée. La deuxième partie de cette thèse a consisté en l'élaboration d'un outil pour évaluer l'état de la santé des populations de tortues d'eau douce sauvages, ce qui reflète également le niveau de contamination des cours d'eau. / Human imprint on Earth is actually so profound leading global environmental changes that affects and modifies the functioning of ecosystems. Among the natural biomes, freshwater ecosystems are remarkable habitats that comprise great species biodiversity but are also the most threatened by human activities. The main causes of the loss of water quality anddisruption of freshwater ecosystems includes water pollution and the introduction of alien species. The fisrt part of this thesis showed invasion of parasites on native freshwater turtle populations, carried by introduced turtle species. Furthermore, it highlighted the necessary to study and describe the new invasive parasite species with more performant technics. The second part of this thesis consisted in the development of a tool to evaluate the status of thehealth of wild freshwater turtle populations, allowed also the level of contamination of the watercourses.

Le commerce mondial et les mouvements humains accroissent les probabilités de transport à longue distance de propagules, et leur introduction dans de nouvelles aires géographiques. Dans certains cas, des espèces récemment établies peuvent devenir dominantes dans la communauté envahie. Malgré les menaces sur les communautés natives et le fonctionnement des écosystèmes, les invasions biologiques constituent des expériences naturelles qui permettent d’étudier les processus éco-évolutifs en temps réel, notamment l’impact de nouvelles interactions biotiques sur la composition et la dynamique des communautés, l’adaptation rapide à de nouvelles conditions environnementales, ou la dispersion en limite de répartition. Les îles océaniques sont particulièrement sensibles aux invasions biologiques en raison de la faible diversité de leurs communautés natives. Dans les terres australes françaises, le carabique marcheur Merizodus soledadinus, natif de Patagonie, a été accidentellement introduit à Kerguelen en 1913. La présente étude vise à comprendre les principaux mécanismes à l’origine du succès invasif de cet insecte aux Iles Kerguelen. Un large ensemble de méthodes ont été utilisée pour explorer les traits écologiques de M. soledadinus, des populations à la molécule. Les analyses génétiques confortent l’hypothèse historique d’un unique évènement d’introduction dans un seul site des Iles Kerguelen. Les populations échantillonnées le long du gradient d’invasion ne montrent pas de structuration génétique. Les traits phénotypiques mesurés montrent une forte différentiation entre les individus selon le temps de résidence des populations, confirmant l’hypothèse de tri spatial des populations au cours de l’expansion géographique. Nous avons démontré que l’expansion géographique et la sélection d’habitats par M. soledadinus est principalement gouvernée par la disponibilité en eau, comme le suggère par la forte sensibilité des adultes au stress hydrique. En parallèle, la colonisation d’habitats en altitude dépend des conditions thermiques, qui semblent être contraignantes pour cet insecte à partir de 200m d’altitude. La colonisation d’habitats d’altitude progresse pourtant, probablement assistée par le changement climatique. Pour finir, les adultes M. soledadinus sont longévifs et actifs toute l’année. Les connaissances apportées sur l’écologie de M. soledadinus et sur la dynamique de son expansion géographique suggèrent la poursuite de la colonisation de l’archipel par ce prédateur. L’ensemble de ces connaissances pourraient être utiles à la paramétrisation d’un modèle d’expansion géographique, qui permettrait de définir les routes de dispersion et les taux d’expansion, dans l’objectif d’assister les mesures de gestion par les agents de la Réserve naturelle des Terres Australes et Antarctiques Françaises. / Global trade and human movements increase the likelihood of long-distance transportation of propagules and their subsequent introduction into new geographic regions. In some instances, newly established species can become dominant in invaded communities, at the expense of native species. Besides threatening invaded communities and ecosystem functions, biological invasions constitute natural experiments that allow to study eco-evolutionary processes in real time, including the occurrence of new biotic interactions affecting community composition, rapid adaptation to novel environmental conditions, or dispersal evolution at range margins. Because of their impoverished native communities, oceanic islands’ ecosystems are particularly sensitive to biological invasions, and the French subantarctic islands are no exception. For instance, the flightless predatory carabid beetle Merizodus soledadinus is native from the southern tip of South America, and has been accidentally introduced to the Kerguelen Islands in 1913. In the present work, we aimed at understanding the main mechanisms underlying the invasive success of this insect at the Kerguelen Islands. Using a vast array of methodologies, ecological features of M. soledadinus were investigated with analytical procedures scaling from population to molecule through the individual level. Genetic investigations support the historically-based hypothesis of a single introduction event at a unique location of the Kerguelen Islands. No genetic structure was observed among individuals sampled from different populations along the invasion gradient. We tested the hypothesis of spatial sorting of populations during range expansion, by exploring phenotypic changes among individuals sampled along the invasion gradient. The measured phenotypic traits revealed major differentiation of adults according to the residence time of their populations, confirming the occurrence of spatial sorting of populations during geographic expansion. We also demonstrated that the geographic expansion of M. soledadinus, and microhabitat selection, are primarily governed by the availability of water resources, as suggested by the high sensitivity to water stress of adults of this ground beetle. In parallel, colonisation of altitudinal habitats is governed by thermal conditions, which seem to be physiologically constraining from 200m asl onwards. As the altitudinal distribution of M. soledadinus still extends, we concluded that ongoing climatic changes play a pivotal role in this expansion. Finally, adults of this ground beetle are long-lived and active year-round. The ecological knowledge of M. soledadinus characteristics and spatial expansion dynamics suggest that the colonisation process of the Kerguelen archipelago by this species will continue. Altogether, these data could be used for parametrising range expansion models that would delineate dispersal pathways and expansion rates, in the objective to assist stakeholders’ management decisions.

Thesis (MSc)--Stellenbosch University, 2013. / ENGLISH ABSTRACT: Successful biological invasions take place when species introduced to regions outside their natural dispersal range overcome several barriers and establish, persist, proliferate and spread potentially resulting in major threats to biodiversity. The success of invasive alien plants depends on species-specific traits and characteristics of the introduced environment. In this thesis I explore which species traits are important and which environmental barriers need to be overcome for an invasion to occur using Proteaceae as a test case. Firstly, I assessed the global introduction history and invasion ecology of Proteaceae - a large plant family with many taxa that have been widely disseminated by humans, but with few known invaders. This revealed that at least 402 species (i.e. 24% of 1674 species in this family) are known to have been moved by humans out of their native ranges, 58 species (14%) have become naturalized and 8 species (2%) are invasive. The probability of naturalization was greatest for species with large native range sizes, low susceptibility to Phytophthora root-rot disease, larger seeds, mammal-dispersed seeds and those with the capacity to resprout after fire or other disturbances. The probability of naturalized species becoming invasive was greater for species with larger range sizes, species used as barrier plants, taller species, species with smaller seeds, serotinous species, and those that regenerated mainly through re-seeding. Secondly, I looked at mechanisms underlying naturalization on a regional scale, using species which are not already classified as major invaders. At least 26 non-native Proteaceae species have been introduced to, and are cultivated in, South Africa. Propagule pressure facilitated the naturalization of Hakea salicifolia populations in climatically suitable areas, but in suboptimal climates human-mediated land disturbance and land management activities are important for naturalization. Similar drivers are important for naturalization of other alien Proteaceae: a long residence time, fire regimes, poor land management, and propagule pressure were important mechanisms for naturalization. Thirdly, I determined whether reproduction, which in part drives propagule pressure, serves as a barrier for naturalization. I examined several Australian Proteaceae species introduced to South Africa and observed that all species were heavily utilized by native nectar-feeding birds and insects. The five Banksia species that were assessed are self-compatible but four species have a significantly higher reproductive output when pollinators visit inflorescences. Fruit production in H. salicifolia does not differ between naturally-pollinated and autonomously-fertilized flowers. Moreover, no significant difference in fruit production was observed between the five pollination treatments (i.e. natural, pollen-supplementation, autonomous, hand-selfed and hand-crossed treatments) and naturalized and non-naturalized populations. However, pollen limitation was detected in non-naturalized populations which received fewer pollinator visits than naturalized populations. Thus, reproduction limits but is not a fundamental barrier to invasion for H. salicifolia. I conclude that reproductive success of the studied Proteaceae, which is a key barrier determining invasiveness, is not limited by autonomous seed set or mutualisms in the introduced range. In this thesis I highlight biogeographical characteristics, a set of life-history traits and ecological traits as important determinants of invasiveness. These traits are in turn dependent on the stage of invasion. Characteristics of the recipient environment are also important drivers of invasions. This study provides a better understanding of plant invasions in general, but the patterns and processes of invasions highlighted in this thesis will be particularly useful for the current and future management of alien Proteaceae in South Africa and elsewhere, as well as, other species that are adapted to Mediterranean and nutrient poor ecosystems. For example, combining traits of invasiveness and susceptible environments will help to identify which non-native species pose a high risk of becoming invasive (e.g. species with large home ranges and barrier plants) and which conditions in the target area are likely to facilitate or exacerbate invasions (e.g. strong climate match and high propagule pressure). / AFRIKAANSE OPSOMMING: Suksesvolle biologiese indringing vind plaas wanneer ʼn spesie geïntroduseer word in ʼn area buite sy natuurlike verspreidings area, sekere versperrings oorkom, vestig, bly voortbestaan, vermenigvuldig en versprei en potensieel ʼn groot bedreiging inhou vir biodiversiteit. Die sukses van uitheemse indringer plante hang af van spesifieke kenmerke van die spesie en kenmerke van die omgewing waarin dit geïntroduseer word. In hierdie tesis maak ek gebruik van Proteaceae om te ondersoek watter kenmerke is belangrik en watter omgewing versperrings moet oorkom word vir indringing om plaas te vind. Ten eerste assesseer ek die wêreldwye introduksie geskiedenis en indringers ekologie van Proteaceae – ʼn groot plant familie wat wyd gebruik word deur mense, maar met min indringer spesies. Dit het gewys dat mense ten minste 402 spesies (dus 24% van die 1674 spesies in die familie) uit die inheemse areas verskuif het, 58 spesies (14%) genaturaliseer het en 8 spesies (2%) indringers geword het. Die moontlikheid van naturalisasie was die grootste vir spesies met ʼn groot inheemse streek, lae vatbaarheid vir Phytophthora wortelvrot, groter sade, dier verspreide sade en die met ʼn vermoë om weer uit te spruit na ʼn vuur of ander versteuring. Die moontlikheid van genaturaliseerde spesies om indringers te word, was groter vir spesies met groter streek grootte, spesies wat as versperring plante gebruik word, hoër spesies, spesies met kleiner sade, serotiniese spesies, en die wat hoofsaaklik voortbestaan as saadspruiters. Tweedens, het ek gekyk na onderliggende meganismes op ʼn regionale skaal, deur gebruik te maak van spesies wat nie alreeds as belangrike indringers geklassifiseer is nie. Ten minste 26 nie-inheemse Proteaceae spesies is alreeds geïntroduseer en word gekultiveer in Suid Afrika. Propaguul druk fasiliteer die naturalisering van Hakea salicifolia populasies in areas met geskikte klimaat, terwyl in areas met ʼn sub optimale klimaat, versteurings deur mense en grond bestuurs aktiwiteite belangrik is vir naturalisering. Die selfde drywers is belangrik vir die naturalisering van ander uitheemse Proteaceae: lang verblyftyd, vuur bestel, swak land bestuur en propaguul druk. Derdens het ek bepaal of reproduksie, wat gedeeltelik propaguul druk dryf, ʼn versperring is vir naturalisasie. Ek het gekyk na verskeie Australiese Proteaceae spesies wat geïntroduseer is in Suid Afrika, en het gevind dat al die spesies besoek word deur inheemse nektar etende voëls en insekte. Die vyf Banksia spesies wat geassesseer is, kan self bestuif, maar vier van die spesies het ʼn betekenisvolle hoër reproduksie wanneer bloeiwyses deur bestuiwers besoek word. Vrug produksie verskil nie tussen natuurlik bestuifde en self bestuifde blomme in H. salicifolia nie. Verder was daar geen verskil tussen vrug produksie van die vyf bestuiwings behandelinge (naamlik: natuurlik, stuifmeel bygevoeg, self, hand self en hand kruis) en tussen genaturaliseerde en nie genaturaliseerde populasies. Ewenwel, stuifmeel beperking is gevind in nie-genaturaliseerde populasies wat egter ook minder besoeke ontvang het dan die genaturaliseerde populasies. Dus, reproduksie kan die verspreiding beperk maar is nie ʼn fundamentele versperring vir indringing van H. salicifolia nie. My konklusies is dat die reproduktiewe sukses, wat andersins ʼn sleutel versperring is vir indringing, in die bestudeerde Proteaceae nie beperk word deur outonomiese saad produksie of mutualismes in die geïntroduseerde gebied nie. In hierdie tesis beklemtoon ek die biogeografiese karakters, lewens geskiedenis kenmerke en ekologiese kenmerke as belangrike bepalers van indringing. Hierdie kenmerke is op hulle beurt weer afhanklik van die stadium van indringing. Karakters van die ontvangende omgewing is ook belangrike dryfvere van indringing. Hierdie studie verbeter hoe ons plant indringing in die algemeen verstaan, maar die patrone en prosesse van indringing wat beklemtoon word in hierdie tesis sal besonder bruikbaar wees vir huidige en toekomstige bestuur van uitheemse Proteaceae in Suid Afrika en op ander plekke, asook vir ander spesies wat aangepas is tot Mediterreense en nutriënt arm ekosisteme. Byvoorbeeld, die kombinasie van kenmerke van indringing en vatbare omgewings sal help om te identifiseer watter uitheemse spesies ʼn hoë risiko inhou om ʼn indringer te word (byvoorbeeld spesies met ʼn groot streek grootte en versperring spesies) en watter kondisies in die teiken area die waarskynlikste indringing fasiliteer of vererger (byvoorbeeld sterk klimaat ooreenstemming en hoë propaguul druk).