Thesis (MSc)--Stellenbosch University, 2013. / ENGLISH ABSTRACT: Alien invasive species are a global concern not only threating biodiversity, but also negatively impacting regional economies. Consequently research aimed at understanding the invasion process is crucial for management of invasive alien species to reduce these impacts. Identifying accurate natal ranges is the key first step for efficient alien plant management, especially biological control. Such knowledge may assist in selecting host-specific biological control agents and help prevent non-target impact on native species. A case in point is the genus Cardiospermum of which species have been extensively moved around the globe with two species now being possible widespread invaders (C. grandiflorum and C. halicacabum), and a third species (C. corindum) having significant invasion potential. However, in some regions the native statuses of these species are not clear, hampering management. In order to minimize potential non-target impacts it is prudent to determine the relationships between Cardiospermum taxa in unknown native ranges prior to the release of biological control agents. With this thesis I aim to review available literature on the genus Cardiospermum, determine potential spread to suitable habitats globally for selected taxa, resolve unknown native ranges in southern Africa, and investigate rapid speciation of an endemic Namib Desert species of balloon vine. Through exhaustive searches for available literature on the genus Cardiospermum I report on the biology and ecology of selected species within the genus, with special focus on the most widespread species. Specifically, using species distribution modelling I investigate the potential spread of C. halicacabum, C. corindum and C. grandiflorum globally and assess the accuracy with which this can be determined using known native ranges to predict current alien ranges. Results indicate that these species have significant potential to spread, though highlighting that species distribution modelling over-fit predicted suitable ranges when using native range data alone. To resolve uncertain native ranges of selected species within the genus I reconstructed a dated multi-gene phylogeny and chloroplast haplotype network to investigate relationships within and among Cardiospermum species. I conclude that C. halicacabum is non-native in southern Africa due to polyphyletic relationships among accessions, while C. corindum is native due to natural long distance dispersal as indicated by a single monophyletic southern African clade. The close relationship observed between C. corindum and the African species, C. pechuelii, lead to a population-level genetic investigation of these two taxa. For this, I constructed a multi-gene phylogeny, chloroplast haplotype network and conducted population genetic diversity analyses which all indicated that the African arid adapted taxa, C. pechuelii, endemic to the Namib Desert of Namibia, evolved from C. corindum following long distance dispersal from South America. This study firstly provides insights into the ecology and biology of Cardiospermum, warning against further introduction due to potential spread. Secondly I resolved the unknown native statutes of Cardiospermum species in southern Africa and, lastly, illustrated the origin of African C. pechuelii. My research findings have major implications for on-going biological research against C. grandiflorum in South Africa and cautions strongly against the release of already-identified biological control agents due to their potential non-target impacts. / AFRIKAANSE OPSOMMING: Indringerspesies is 'n wêreldwye probleem as gevolg van hul direkte bedreiging van biodiversiteit asook hul negatiewe impakte op streeksekonomieë. Gevolglik is navorsing aangaande die indringingsproses van uiterste belang om doeltreffende beheer van indringerspesies te verseker. Die bepaling van die inheemse herkoms van indringerpopulasies is die eerste belangrike stap tot doeltreffende beheer, veral biologiese beheer. Hierdie informasie kan help met die identifisering van gasheer-spesifieke biologiese agente en om potensiële nie-teiken impakte op inheemse plant spesies verhoed. ‘n Voorbeeld is ballon rankplante in die genus Cardiospermum, waarvan verskeie spesies wêreldwyd verspreid is, met die gevolg dat minstens twee spesies (C. halicacabum en C. grandiflorum) nou moontlike wydverspreide indringers is en ‘n derde spesie (C. corindum) ook potensiaal toon as ‘n indringerspesie. In sommige streke is die in- of uitheemse statusse van hierdie spesies onbekend en belemmer gevolglik hul doeltreffende beheer. Met hierdie tesis streef ek om die genus Cardiospermum te hersien en potensiële verspreiding van geselekteerde spesies na geskikte habitatte te bepaal, om onbekende inheemse streke binne Suider-Afrika op te los en laastens, om die herkoms van ‘n ware Afrika spesie (C. pechuelii) te ondersoek. Na deeglike ondersoek van beskikbare literatuur doen ek verslag omtrent die biologie en ekologie van spesies binne Cardiospermum, met spesiale fokus op die mees wydverspreidste spesies. Deur gebruik te maak van spesies verspreidings modelle bestudeer ek die potensiaal van verdere verspreiding wêreldwyd vir C. halicacabum, C. corindum en C. grandiflorum en rapporteer ook omtrent die akkuraatheid van hierdie tegniek. Resultate toon dat alle spesies aansienlike potensiaal toon om verder te versprei, maar beklemtoon ook dat hierdie metode inheemse streek data “oor-aanpas”. Om enige onsekerhede aangaande inheemse streke op te los, het ek ‘n gedateerde multi-geen filogenie en ‘n chloroplas haplotipe netwerk opgetrek om die verhoudings binne en tussen geselekteerde Cardiospermum spesies te bepaal. Ek kom tot gevolgtrekking dat C. halicacabum nie inheems in Suider-Afrika is nie, as gevolg van polifiletiese verhoudings, en dat C. corindum inheems is en moontlik ook ‘n voorbeeld van natuurlike langafstand verspreiding, soos aangedui deur ‘n enkele monofiletiese groep. Die naverwante filogetiese verhouding tussen C. corindum en C. pechuelii het daartoe gelei dat ek die herkoms van C. pechuelii in Afrika ondersoek het. ‘n Multi-geen filogenie, chloroplas haplotipe netwerk en populasie genetiese diversiteit analises het aangedui dat C. pechuelii endemies is tot die Namib Woestyn van Namibië, en die produk is van lang afstand verspreiding van C. corindum vanaf Suid-Amerika. Hierdie studie gee eerstens insig tot die ekologie en biologie van Cardiospermum en waarsku teen verdere verspreiding. Tweedens dui my studie ook die opgeloste inheemse streke in Suider-Afrika aan van sekere Cardiospermum spesies, en laastens, illustreer ek die herkoms van C. pehuelii. My navorsing het baie belangrike gevolge vir die huidige biologiese beheer program teen C. grandiflorum in Suid-Afrika en waarsku teen die vrystelling van reeds geïndentifiseerde biologiese beheer agente weens potensiële nie-teiken impakte op inheemse spesies.

Thesis (MSc)--Stellenbosch University, 2013. / ENGLISH ABSTRACT: This thesis explores how human dictated methods of introduction and species-specific traits interact to define spatial patterns in invasive plant populations using Acacia elata as a model species. I initially asked whether the relatively small invasive extent (when compared to congeners introduced for forestry or dune stabilization) of a species used widely for ornamental purposes (A. elata) is due to low rates of reproduction in South Africa. Results indicate that A. elata has similar traits to other invasive Australia Acacia species: annual seed input into the leaf litter was high (up to 5000 seeds m-2); large seedbanks develop (>20 000 seeds m-2) in established stands; seed germinability is very high (>90%); seeds accumulate mostly in the top soil layers but can infiltrate to depths of 40cm; and seed germination appears to be stimulated by fire. I argue that the current limited distribution of invasive A. elata populations is not attributable to species-specific reproductive traits. Following on from this I addressed the relative importance of species traits, the recipient environment, and introduction dynamics using species distribution models and spread models defined using a range of parameters representative of invasive Australian acacia species. Results indicate that in the early stages of simulated invasions, the number of trees introduced was the most important parameter influencing abundance and extent, while the placement of introduction foci in urban areas associated with ornamental introductions limited the extent and abundance of invasive populations. I suggest that conditions relating to human-mediated introduction events initially mask the influence of intrinsic species traits and help to explain the failure and success of species associated with specific human-shaped pathways. Third, I used scale-area curves to determine how these human-dictated introduction conditions influenced the spatial structure of populations. The fractal dimension (Dij) of A. elata populations indicates plot scale (2.5-25m) spread with high densities that are increasing. Artefacts of introduction history were observed at the regional scale (2.5-25km) - populations are contiguous around introduction foci such as towns and roads. Moreover, fragmentation of A. elata populations at local to landscape scales (0.25-2.5km) is due to a combination of the haphazard regional placement and limited secondary dispersal vectors, both of which can be linked to their introduction history. In summary, this study used a combination of field-based data, modelling techniques and broad-scale sampling methods to assess how intrinsic species traits and introduction conditions mediate invasive spread. I conclude that introduction history can have a long-lasting (100+ years) influence on the spatial structure and distribution of invasive plants, which can mask the influence of individual species traits. / AFRIKAANSE OPSOMMING: Alhoewel menigte plant spesies deur mense ingevoer word, word slegs 'n klein hoeveelheid van plante indringers. Die beperkte aantal spesies wat in nuwe habitatte vestig en versprei, kan grotendeels toegeskryf word aan die omgewing en mensgemaakte hindernisse ten opsigte van oorlewing, voortplanting en verspreiding. In hierdie tesis gebruik ek Acacia elata as ʼn model spesie om te ondersoek hoe die invoeringsproses sowel as spesie spesifieke eienskappe die ruimtelike struktuur van indringende populasies bepaal. Aanvanklik was my vraag of die relatiewe klein indringingsarea van ʼn spesie wat ingevoer is vir ornamentele doeleindes (Acacia elata), teenoor die groot indringingsareas van Australiese Acacia-spesies wat ingevoer is vir bosbou of duin stabilisering, die gevolg is van lae voortplantings tempo in Suid-Afrika. Resultate dui daarop dat A. elata soortgelyke eienskappe as ander indringende Australië akasia spesies het: jaarlikse saad produksie is hoog (tot 5000 sade m - 2); groot ontwikkelde saadbanke (> 20 000 sade m - 2); hoë saad ontkieming (> 90 %); saad versamel in die boonste grondlae, alhoewel dit kan infiltreer tot 'n diepte van 40 cm; laastens, ontkieming word deur vuur gestimuleer. Hierdie resultate stel voor dat die huidige beperkte verspreiding van indringende A. elata populasies nie die gevolg is van spesie spesifieke voortplantings eienskappe nie. Gevolglik het ek 11 indringende Australiese akasia-spesies ondersoek om die relatiewe belangrikheid van spesie eienskappe, die area waar hul geplant is, asook invoerings aspekte te ondersoek. Resultate wys dat die aantal bome wat ingevoer is, die belangrikste eienskap was ten opsigte van digtheid en verspreiding van ʼn indringende spesie, terwyl invoering naby fokuspunte in stedelike gebiede weer spesie digtheid en verspreiding beperk. Hierdie resultate toon dat spesies eienskappe aanvanklik nie ʼn groot invloed het op hul sukses nie, maar dat hul sukses eerder afhang van waar hul geplant word. Derdens, gebruik ek skaal - area kurwes om te bepaal hoe die invoering van A. elata die ruimtelike struktuur van populasies beïnvloed het. Die fraktale dimensie (Dij) van A. elata bevolkings dui op verspreiding by die plaaslike vlak (2,5 - 25m), wat voorstel dat verspreiding nie op hierdie skaal beperk is nie. Eienskappe van die rede/metode van invoering was wel waargeneem op die groter skaal (2.5 - 25km) waar populasies digter was naby dorpe en paaie. Die intermediêre skaal (0,25 - 2.5km) dui daarop dat A. elata populasies huidiglik beperk word deur 'n kombinasie van lukrake plasing en beperkte sekondêre verspreiding, albei wat gekoppel kan word aan hul invoer geskiedenis. Hierdie studie maak dus gebruik van 'n kombinasie van veld gebaseerde data, modellerings tegnieke en breë skaal steekproefmetodes om te evalueer hoe algemene spesies eienskappe en invoer toestande verspreiding bepaal. Sodoende verskaf ek ʼn raamwerk om die invloed van invoer toestande op die ruimtelike struktuur en verspreiding van ornamentele plant indringers te verstaan.

Thesis (MSc)--Stellenbosch University, 2013. / ENGLISH ABSTRACT: The generally low concentrations of P and N in the soil, causes most plants to experience nutrient deficiency during their life cycle. Lupins can rely on both cluster roots and nodules for P acquisition and biological nitrogen fixation (BNF) respectively. The legume Lupinus albus is able to survive under low nutrient conditions, because it has two specialized belowground organs for the acquisition of N and P. In this regard, cluster roots increase P uptake and root nodules acquire atmospheric N2 via biological nitrogen fixation (BNF). Although these organs normally tolerates low P conditions, very little is known about their physiological and metabolic flexibility during variations in P supply. Furthermore, the resource allocation (C, N and P) between cluster roots and nodules has also been largely understudied. The aim of this investigation was therefore to determine the resource allocation, physiological and metabolic flexibility of these organs during variations in P supply. Although variation on P supply had no effect on the total biomass, there were significant differences in specialised below-ground organ allocation to cluster roots and nodule formation. Cluster root formation and the associated C-costs increased during low P supply. In contrast to the cluster root decline at high P supply, there was an increase in nodule growth allocation and corresponding C-costs. Since cluster roots were able to increase P acquisition under low P conditions, this below-ground investment may also have benefited the P nutrition of nodules. These findings provide evidence that when lupins acquire N via BNF in their nodules, there may be a trade-off in resource allocation between cluster roots and nodules. The short-term elevated P supply, caused an increased allocation of C and respiratory costs to nodules, at the expense of cluster roots. This alteration was also reflected in the increase in nodule enzyme activities related to organic acid synthesis, such as Phosphoenol-pyruvate Carboxylase (PEPC), Pyruvate Kinase (PK), Malate Dehydrogenase (NADH-MDH) and Malic Enzyme (ME). In cluster roots, the elevated P conditions, caused a decline in these organic acid synthesizing enzymes. This suggests that during short-term elevated P supply, there is a great degree of physiological and metabolic flexibility in the lupin nutrient acquiring structures. / AFRIKAANSE OPSOMMING: Die algemeen lae konsentrasies van fosfaat en stikstof in die grond , veroorsaak dat die meeste plante voedingstekorte ervaar tydens hul lewensiklus. Lupiene kan staatmaak op beide groep-wortels en wortel-knoppies vir P verkryging en biologiese stikstofbinding onderskeidelik. Die peulplant Lupinus albus is in staat om te oorleef onder lae voedings toestande , as gevolg van die twee gespesialiseerde ondergrondse organe vir die verkryging van stikstof en fosfaat. In hierdie verband verhoog groep-wortels fosfaat opname en wortel-knoppies verkry atmosferiese stikstof via biologiese stikstofbinding. Alhoewel hierdie organe normaalweg lae fosfaat toestande verdra , is baie min bekend oor hul fisiologiese en metaboliese buigsaamheid tydens variasies in fosfaat aanwending. Daar is verder ‘n tekort aan die studie van hulpbron toekenning tussen groep-wortels en wortel-knoppies. Die doel van hierdie ondersoek was dus om die toekenning van hulpbronne , fisiologiese en metaboliese buigsaamheid van hierdie organe tydens variasies in fosfaat aanwending te bepaal. Variesie in fosfaat verskaffing het geen invloed op die totale plant biomassa gehad nie, maar daar was wel ‘n beduidende verskil in gespesialiseerde ondergrondse toekenning tussen groep- wortels en wortel-knoppies. Groep-wortel vorming en die gepaardgaande koolstof koste het toegeneem met lae fosfaat verskaffing. In teenstelling met die groep-wortel daling met hoë fosfaat verskaffing, was daar 'n toename in groei van wortel-knoppies en die ooreenstemmende koolstof koste daarvan. Aangesien groep-wortels in staat was om fosfaat verkryging te verhoog onder lae fosfaat toestande, mag hierdie ondergrondse belegging bygedra het tot die voeding van wortel-knoppies . Hierdie bevindings bewys dat lupiene afhanklik van wortel-knoppies ‘n wisselwerking in toekenning van hulpbronne, tussen groep-wrotels en wortel-knoppies handaaf. Kort termyn verhoogde fosfaat aanwending veroorsaak 'n verhoogde toekenning van koolstof en respiratoriese energie na wortel-knoppiess, ten koste van groep-wortels . Hierdie verandering is ook weerspieël in die toename in wortel-knoppie ensiem aktiwiteit in verband met organiese suur sintese (PEPC PK,MDH,ME) . In groep-wortels, het die verhoogde P toestande verder 'n afname in die organiese suur produserende ensieme veroorsaak. Dit dui aan dat tydens kort termyn verhoogde P aanwending, daar 'n groot mate van fisiologiese en metaboliese buigsaamheid in die lupiene voedingstowwe verkryging strukture plaasvind.

Thesis (PhD (Botany and Zoology))—University of Stellenbosch, 2009. / The genus Oxalis forms a major part of the flora of southern Africa, in particular the Cape Floristic Region (CFR) at the southwestern tip of the continent, but the current taxonomy is outdated and ecological knowledge of the lineage is sadly incomplete. In this thesis I set out to address several aspects of Oxalis systematics that urgently require attention. Firstly, the current macro-morphological taxonomy requires phylogenetic testing, as it is acknowledged to be incomplete and artificial. I address this need by providing a DNA sequence-based phylogeny of three markers, using three different inference methods, for nearly three quarters of the indigenous species. This phylogeny confirmed both the monophyly of the southern African taxa, and the artificiality of the current classification system. It is congruent with previous sequence-based reconstructions of smaller groups of southern African Oxalis species, and with the palynological classification proposed for the genus. Secondly, previous phylogenetic work on the southern African members could not resolve basal relationships within the southern African clade. I attempt to address this problem by sequencing three extra chloroplast markers for a select group of taxa, followed by separate and combined (total evidence) molecular phylogenetic analyses. This approach did increase resolution at the base of the southern African lineage, but many clades still showed poor resolution and support despite the use of more than 7 000 bases of sequence data. Resolving these clades within the southern African Oxalis phylogeny remains a challenge, and should prove a fertile field for future research. Thirdly, the ages (and thus duration of presence) of many Cape plant lineages within the CFR are of major interest, given that the CFR represents a global biodiversity hotspot. The age of the genus in the Cape is estimated by analyzing combined sequence data for all sampled taxa under both a Bayesian Relaxed Clock and a semiparametric Penalised Likelihood method, using calibration points inferred from Relaxed Clock analyses of the entire order Oxalidales, for which fossil data are available. In an attempt to account for known problems with divergence time estimation, I explored the potential bias introduced by method used, marker genome source and different calibrations on the root. The results indicate substantial variation in the age of crown southern African Oxalis over a nearly twenty million year period, varying according to source data, calibration estimate and methodology employed in the reconstruction. Despite this major variability, all average estimates are older than iv 18 million years, which agrees with a growing body of evidence that there has been a gradual accumulation of floristic diversity in the CFR, rather than a rapid, recent burst of speciation. Fourthly, as the produced phylogenies conclusively show the artificial nature of the current taxonomy, I propose a new, almost completely different classification for southern African Oxalis taxa. Although a significant improvement, this classification is considered informal due to the complete disagreement between the old and proposed new taxonomies, poor resolution in some of the proposed lineages, and a need to confirm proposed groups (clades) with the identification of morphological synapomorphies. Potential synapomorphies for various clades are proposed and discussed, which should guide future research. Fifthly, the presence of bulbs in this genus is of great interest as a potential preadaptation for seasonally arid climates. The evolution of the bulbous habit in Oxalis is here explored for the first time. I address the sequence of major morphological character state changes leading to the suite of characters corresponding to the bulbous habit. The homology of basal leaf petioles, fleshy leaf scales and tunics is discussed, and it is shown that many bulb characters present in the southern African lineage are also found in the close relatives of this lineage, and are thus older than this lineage. The ecological and evolutionary implications of bulb geophytism in the CFR Oxalis are also discussed. Finally, I address certain taxonomic issues that arose during the course of this study. Co-authors and I describe the new species O. saltusbelli and O. ericifolia. We also clarify issues surrounding the tremendously variable group species O. flava and propose some nomenclatural changes and synonyms for related taxa. We also address the taxonomic position of the rare species O. purpurata, which was located too late in the course of this study to include in the main analyses.

Thesis (PhD (Botany and Zoology))—-Stellenbosch University, 2008. / Streptocarpus Lindl. (Gesneriaceae DC.) is a genus of herbaceous plants containing approximately 160 species, of which the majority occur in Africa and Madagascar. They are largely restricted to shaded and moist habitats such as primary forest and rock outcrops. The genus contains considerable morphological variation, with subgenus Streptocarpella containing caulescent species and subgenus Streptocarpus mostly consisting of acaulescent growth forms, mainly the unifoliates, plurifoliates and rosulates. Preliminary molecular analyses conducted using nuclear ITS sequence data suggested that subgenus Streptocarpus evolved in tropical central Africa, before radiating in several independent waves into southern Africa. Streptocarpus has therefore only recently spread into South Africa. Amongst the South African species, 11 morphologically similar species were identified as being closely related, together forming the Cape primrose clade, based on the analysis of nuclear ITS sequence data. However, these analyses only contained a few South African species, and the ITS data did not provide enough resolution of relationships within this clade. In this study nuclear and chloroplast sequence data as well as nuclear microsatellite data were therefore employed to unravel the complex relationships amongst the South African Streptocarpus species. The analyses indicate that 16 rosulate, palynologically similar species (S. primulifolius, S. rexii, S. johannis, S. baudertii, S. modestus, S. formosus, S. gardenii, S. lilliputana, the S. cyaneus complex [S. cyaneus, S. parviflorus, S. fenestra-dei, S. kunhardtii and S. roseo-albus], S. floribundus, S. aylae and S. kentaniensis), the core Cape primrose species, are closely related, while five unifoliate/plurifoliate, palynologically more variable species (S. denticulatus, S. dunnii, S. pusillus, S. rimicola and S. bolusii) consistently emerged as more distantly related to the core Cape primrose species. However, the positions of a further ten species (S. meyeri, S. montigena, S. fanniniae, S. caeruleus, S. longiflorus, S. polyanthus, S. saundersii, S. porphyrostachys, S. grandis and S. vandeleurii) were more complex in the analyses, indicating that hybridization has played a role in their evolution. Five of these species (S. meyeri, S. montigena, S. fanniniae, S. caeruleus and S. longiflorus) are, however, palynologically homogenous and rosulates, and therefore are probably more closely related to the core Cape primrose species, while the other five (S. polyanthus, S. saundersii, S. porphyrostachys, S. grandis and S. vandeleurii) are unifoliates/plurifoliates that are palynologically more heterogenous, and are probably more distantly related to the core Cape primrose species. Amongst the core Cape primrose species, S. primulifolius emerged as being ancestral or having hybridized with many of the other species, while the S. cyaneus complex forms a geographically and genetically more isolated group. However, evolutionary relationships amongst these species were to a certain extent obscured by incomplete lineage sorting caused by limited interpopulation gene flow, frequent hybridization and rapid speciation. The analyses confirmed that the Pondoland Centre forests constitute important Pleistocene refugia, and revealed some of the historical migration routes along which the species had radiated.

Thesis (MSc (Botany and Zoology))--Stellenbosch University, 2011. / ENGLISH ABSTRACT: See full text for abstract / AFRIKAANSE OPSOMMING: sien volteks vir opsomming

Thesis (MSc)--University of Stellenbosch, 2004. / ENGLISH ABSTRACT: Arbuscular mycorrhizal (AM) C-costs in grapevines were investigated. Since both dormant vines and AM colonisation rely on stored C for initial growth, AM colonisation costs would therefore compete with plant growth for available C reserves. The aims of this study were to assess the host C economy during AM development and the subsequent C-costs of N and P uptake, as well as the effects of C costs on host growth. This was evaluated in two separate experiments; one assessing the symbiotic influence on the C costs of fungal establishment and nutritional benefits, whilst the other one evaluated the effects of the symbiosis on host growth and nutrient productivities. This study has shown that AM acts as a C sink, competing with the host for available C. Past work on the AM sink effect has focused mainly on the movement of photosynthetic C below ground to support the AM fungus. This however, does not take into account the effect that stored C will have on the C economy of the plant and symbiosis. The role of stored C becomes even more crucial when working with deciduous plants that rely on stored C for new growth at start of a growing season. It has been reported that stored C in AM plants is remobilized at the start of a growing season and then the C reserves are refilled towards the end of the season, when the plants enter dormancy. The initial costs of AM fungal colonisation were borne by the above-ground C reserves, at the expense of new growth in host plants. These costs were offset once the plateau phase was reached, and the depleted reserves started to refill. Once established, the active symbiosis imposed a considerable below ground C sink on host reserves. In spite of these costs, the improved P nutrition of AM roots was achieved with a more efficient C-use. This concurs with other findings, that of the belowground C allocated to AM roots, a greater part is used by AM respiration and a smaller part for P uptake. The C costs of the AM fungal phase of rapid development can be seen as negative to root growth and shoot development. These negative effects may continue for a period of time, even during the plateau phase of fungal development. Once the AM symbiosis is fully established, the host growth and development is then improved to a greater extent than in non-AM plants. From this study it can be concluded that AM growth directly competes with host development, but the symbionts revert to a beneficial partnership once it is fully established. / AFRIKAANSE OPSOMMING: Die C koste van arbuskulêre mikorisa (AM) in wingerdstokke is ondersoek. Beide rustende wingerdstokke en AM koloniseering is afhanklik van gestoorde C vir aanvanklike groei. AM kolonisering sou dus met plantgroei kompeteer vir beskikbare C reserwes. Die doelstellings van hierdie ondersoek was eerstens om die C-ekonomie van die gasheer tydens AM ontwikkeling en die gevolglike Ckostes van N en P opname te bepaal en tweedens sowel as die invloed van C veranderings op gasheergroei vas te stel. Hierdie is in twee afsonderlike eksperimente ondersoek: een om die simbiotiese invloed op die C-kostes van swam-vestiging en voedingsvoordele te bepaal, terwyl die ander die uitwerking van simbiose op gasheergroei en voedings doeltreffenheid evalueer het. Die ondersoek het bewys dat AM, as ‘n C-sink, kompeteer met die gasheer vir beskikbare C. Vorige werk oor die AM sink-effek het hoofsaaklik gefokus op die afwaartse beweging van fotosintetiese C om die AM-swam ondergronds te ondersteun. Die werk neem egter nie in ag wat die effek van gestoorde C op die C-ekonomie van die plant en simbiose sou wees nie. Die rol van gestoorde C is selfs nog meer belangrik wanneer met bladwisselende plante gewerk word, omdat sulke plante op gestoorde-C vir nuwe groei aan die begin van die groeiseisoen staatmaak. Dit is op rekord dat gestoorde C in bladwisselende plante by aanvang van die groeiseisoen gemobiliseer word en dat die C-reserwes teen die einde van die seisoen wanneer die plante rustyd nader, weer hervul word. Die aanvanklike kostes van AM kolonisering is deur die bogronds C-reserwes, ten koste van nuwe groei van die gasheerplante, gedra. Hierdie kostes herstel sodra die plato-fase bereik is, waar die uitgeputte reserwes begin hervul het. As die aktiewe simbiose eers gevestig is, sal dit as ‘n onderg P-voeding van AM wortels verkry wordrondse C-sink vir gasheer optree.Hierdie C verbruik word egter as doeltreffend beskou aangesien verbeterde. Dit is bekend dat ‘n groter deel van die ondergrondse C geallokeer word aan AM-wortels, deur middel van AM respirasie en P-opname. Die C-kostes van die AM-fungus tydens die fase van vinnige ontwikkeling, kan ‘n negatiewe effek op wortel- en lootontwikkeling hê. Hierdie negatiewe uitwerking kan vir ‘n tydperk voortdeur, selfs gedurende die plato-fase van fungi-ontwikkeling. Sodra die AM-simbiose volledig gevestig is, word gasheergroei en ontwikkeling tot ‘n groter mate verbeter as in plante sonder AM-fungi. Hierdie ondersoek het bewys dat AM groei direk met gasheerontwikkeling kompeteer, maar dat die simbiose ‘n voordelige vennootskap vorm sodra dit volledig gevestig is.

Thesis (MSc)--Stellenbosch University, 2012. / ENGLISH ABSTRACT: This thesis investigates the invasiveness and current status of two Acacia species recently identified as invaders in South Africa in order to determine the feasibility of their eradication. Australian acacias are among South Africa’s worst invasive species and many have had widespread damaging impacts on native ecosystems. In addition, several Acacia species still exist as small isolated populations in the country and have been targeted for eradication in order to prevent potential widespread impacts. This work assesses Acacia implexa (Chapter 2) and Acacia stricta (Chapter 3) as potential eradication targets by quantifying the extent of their invasion in South Africa, assessing the risk they pose to the country and evaluating the feasibility of their eradication based on estimated costs of clearing. Results of formal risk assessments show that both A. implexa and A. stricta should be considered high risk species, and bioclimatic model predictions indicate that both species have large potential ranges in South Africa. Detailed population surveys found that A. implexa and A. stricta each occur at several distinct localities all in the Western Cape Province. Acacia implexa populations were found at three sites (Tokai, Wolseley and Stellenbosch) where they have densified by means of vegetative suckering allowing A. implexa to outcompete native vegetation. No evidence of large seed banks of A. implexa were found, however vigorous resprouting following damage makes the control of A. implexa difficult. Acacia stricta was found at nine localities all in the Knysna area of the Garden Route, where populations are spreading along disturbed roadsides in plantations. Acacia stricta produces large amounts of seeds and can accumulate large seed banks. Seed spread is most likely due to large-scale soil movement by road maintenance vehicles which can easily lead to the establishment of new populations. We therefore used a predictive risk mapping approach based on the association of A. stricta to roadsides and disturbed plantations to enable effective searching to detect all infestations of A. stricta. Based on the high risk of both species and the limited range sizes of the currently known populations, we recommend that A. implexa and A. stricta remain targets for eradication. Management strategies proposed for these species (Chapter 4) include clearing on an annual (in the case of A. stricta) or biannual (for A. implexa) basis to prevent seed production, and targeted awareness campaigns at a national scale to determine whether our current knowledge of the extents of A. implexa and A. stricta are accurate. This work has shown that detailed assessments of species at intermediate stages of invasion is an important initial step in an eradication attempt, and better understanding of species specific invasion characteristics can help to improve management and potentially increase the probability of success of eradication. / AFRIKAANSE OPSOMMING: Hierdie tesis ondersoek die invasieve en die huidige status van twee Acacia spesies onlangs geïdentifiseer as indringers in Suid-Afrika ten einde die lewensvatbaarheid van hul uitwissing om te bepaal. Australiese akasias is onder Suid-Afrika se ergste indringerspesies en baie het wydverspreide skadelike impak op die inheemse ekosisteme. Verder het verskeie Acacia spesies bestaan nog steeds as 'n klein geïsoleerde bevolkings in die land en wat geteiken is vir uitwissing in om moontlike grootskaalse impakte te voorkom. Hierdie werk beoordeel Acacia implexa (Hoofstuk 2) en Acacia stricta (Hoofstuk 3) as 'n moontlike uitwissing teikens deur die kwantifisering van die omvang van hul inval in Suid-Afrika, die beoordeling van die risiko wat hulle inhou vir die land en die evaluering van die haalbaarheid van hul uitwissing op grond van beraamde koste van die wiele ry. Resultate van formele risikobepalings toon dat beide die A. implexa en A. stricta moet oorweeg word om 'n hoë risiko spesies, en bioclimatic model voorspellings dui daarop dat beide spesies het 'n groot potensiaal bereik in Suid-Afrika. Uitgebreide bevolkings opname gevind dat A. implexa en A. stricta elk by verskeie afsonderlike plekke in die Wes-Kaap voorkom. Acacia implexa is op drie plekke (Tokai, Wolseley en Stellenbosch) gevind, waar hulle deur middel van vegetatiewe suier densified en inheemse plantegroei oorwin het. Geen bewyse van groot nageslag banke van A. implexa is gevind, maar in kragtige resprouting volgende skade maak die beheer A. implexa moeilik is. Die Acacia stricta is op nege plekke in die Knysna-omgewing van die Tuinroete, waar die bevolkings verspreiding langs die versteurde paaie in plantasies. Acacia stricta produseer groot hoeveelhede saad en kan versamel groot saadbanke. Saad versprei is waarskynlik te danke aan grootskaalse grond beweging deur die instandhouding van paaie voertuie wat kan lei tot die vestiging van nuwe bevolkings. Ons het dus 'n voorspellende risiko kartering benadering wat gebaseer is op die vereniging van A. stricta aan paaie en versteurde plantasies in staat te stel om doeltreffend te soek alle besmettings van A. stricta op te spoor. Gegrond op die hoë risiko van beide spesies en die beperkte reeks groottes van die bevolking wat tans bekend is, beveel ons aan dat A. implexa en A. stricta bly teikens vir uitwissing. Bestuurstrategieë vir hierdie spesies (Hoofstuk 4) voorgestel word, sluit in die skoonmaak op 'n jaarlikse (in die geval van A. stricta) of die halfjaarlikse (vir A. implexa) basis van die saad produksie, en geteikende bewusmakingsveldtogte om te voorkom dat 'n nasionale skaal om te bepaal of ons huidige kennis van die omvang van A. implexa en A. stricta akkuraat is. Hierdie werk het getoon dat uitgebreide aanslae van spesies op intermediêre fases van die inval is 'n belangrike eerste stap in 'n poging van die uitwissing, en 'n beter begrip van spesies spesifieke inval eienskappe kan jou help om te verbeter en potensieel verhoog die waarskynlikheid van sukses van die uitroeiing nie.

Thesis (MSc (Botany and Zoology))--University of Stellenbosch, 2010. / ENGLISH ABSTRACT: It is thought that a large proportion of the great variety of floral structures in flowering plants reflect adaptations to different biotic pollen vectors. Divergence in flower traits and pollinators is linked to speciation. Pollinator-driven speciation is thought to have played a large role in the spectacular floral diversity found in South African Iridaceae and the genus Tritoniopsis is a particularly good example of this. This study focuses on Tritoniopsis revoluta, a pink irid occurring in the Swartberg and Langeberg Mountains, as well as Potberg Mountain. I tested the hypothesis that variation in flower tube-lengths of Tritoniopsis revoluta are related to the geographic distribution of pollinators and the variation of their tongue lengths. It was determined that this species is highly variable in respect to corolla tube-length and is pollinated by different fly species across its range. Also, the tongue-lengths of the fly pollinators corresponded almost exactly with the tube-length of the flowers they were pollinating in each population. In some populations, where long-proboscid flies were absent, bees were observed visiting T. revoluta flowers. This presents evidence for pollinator-driven floral variation within a single plant species, and most of this vast diversification in floral morphology has probably been driven by morphological variation found within a single fly family. In one population I found variable tube-lengths which appeared to exhibit a bimodal distribution of corolla tubelengths. I hypothesized that the two Tritoniopsis revoluta ecotypes at this population are pollinated by two different pollinators, leading to assortative mating, and ultimately strong inter-ecotype incompatibility. Tritoniopsis revoluta is self-incompatible and exists as two discrete entities (morphotypes) at the Gysmanshoek Pass site, and these entities differ in tubelength, color, nectar volume and sugar content. These morphotypes were not pollinated by long-proboscid flies, but seems to represent a recent shift to pollination by Amegilla bees. However, ecotypes are not reproductively isolated as short and long flowers can produce offspring, rather tube-length differences are possibly maintained through spatial separation. To compliment the correlatory data between flower tube-lengths and pollinator tongue-lengths, I used molecular tools (chloroplast markers and AFLPs) to elucidate the patterns of tube-length evolution in Tritoniopsis revoluta. I aimed to determine the directionality and frequency of transitions between tube-length categories. Tube-length transitions would be suggestive of flower morphology being labile, and together with the tube-tongue length correlation it suggests pollinator shifts may drive the changes in tube length. Character state reconstructions using tube-length as character determined that four evolutionary transitions to shorter tube-length categories and two transitions to longer categories occurred. I also tested whether morphological divergence between populations corresponds to patterns of divergence from neutral genetic markers. Population genetic structure in this system showed that the different populations of T. revoluta are vicariant and tube-length differences between them could have evolved through selection. / AFRIKAANSE OPSOMMING: Dit is ‘n algemene gedagte dat die groot verskeidenheid blom strukture in die angiosperme dui op aanpassings tot verskillende biotiese stuifmeel draers. Die diverse blom strukture in baie van die groot Kaapse genera kan verduidelik word deur aanpassings tot veranderinge in bestuiwings-sisteme. ‘n Aantal studies hieroor stel voor dat bestuiwers nie net die veranderinge in blom morfologie bewerkstellig nie, maar ook ‘n rol speel in die aanpassende uiteenlopendheid van blomplant kenmerke. Spesiasie bewerkstellig deur bestuiwers het moontlik ‘n groot rol gespeel in die blom-diversiteit wat gevind word in die Suid-Afrikaanse Iridaceae familie, en die genus Tritoniopsis is ‘n baie goeie voorbeeld hiervan. Hierdie studie fokus spesifiek op Tritoniopsis revoluta, ‘n pienk iris wat voorkom in die Swart- en Langeberge, asook by Potberg. Die hipotese dat die variasie in buis-lengtes van T. revoluta verwant is aan die geografiese verspreiding van bestuiwers en die variasie in hul tong-lengtes is hier getoets. Dit is bepaal dat hierdie spesie groot variasie toon in terme van buis-lengtes en bestuif word deur verskillende vlieg spesies regoor sy verspreiding. Die tong-lengtes van hierdie vlieë korrespondeer ook met die buis-lengtes van die blomme wat hul bestuif in elkeen van die T. revoluta populasies. In sommige van die populasies, waar lang-tong vlieë afwesig was, is bye wat die T. revoluta blomme besoek, opgemerk. Hierdie resultate lewer bewyse vir die hipotese dat bestuiwers blom morfologie kan beïnvloed; die interessante hiervan is dat die variasie in buis-lengtes in hierdie spesie heel moontlik te danke is aan die morfologies variasie wat gevind word in ‘n enkele lang-tong vlieg familie. In een van die populasies het ek ‘n bimodale verspreiding van buis-lengtes gevind. ‘n Logiese afleiding is dat hierdie twee verskillende buislengtes – ekotipes – deur twee verskillende bestuiwers besoek word, en dat dit lei tot sterk onversoenbaarheid tussen ekotipes. Tritoniopsis revoluta is nie instaat tot self-bestuiwing nie en die twee ekotipes verskil in terme van buis-lengtes, kleur, nektar volume en suiker inhoud. Kort- en lang-buis blomme word nie eksklusief bestuif deur lang-tong vlieë in die Gysmanshoek Pas nie, maar word in die algemeen ook bestuif deur bye van die genus Amegilla. Die twee ekotipes is in staat om te reproduseer met mekaar, so die buis-lengte verskille word moontlik in stand gehou deur hul geografiese skeiding. Om die korrelasie analise tussen blom buislengtes en vlieg tong-lengtes te komplimenteer, het ek molekulêre tegnieke (chloroplast merkers en AFLPs) gebruik om die patrone van buis-lengte evolusie in Tritoniopsis revoluta duidelik te maak. Ten eerste het ek bepaal of verkortings en verlengings van buis-lengtes een keer in die verlede gebeur het, of as meermalige gebeurtenisse. Meermalige veranderinge in buis-lengtes kan moontlik dui op verskuiwings tussen verskillende bestuiwers, asook taksonomiese verdelings wat korrespondeer met bestuiwer veranderinge. Ek het ook bepaal of die buis-lengte verskille in die verskillende populasies toegeskryf kan word aan seleksie prosesse. Deur buis-lengte as karakter te gebruik, het ek karakter-status rekonstruksies gedoen en bepaal dat vier ewolutionêre transisies na korter buis-lengte kategorieë, en twee transisies na langer kategorieë plaasgevind het. Populasie genetiese struktuur in die sisteem dui daarop dat T. revoluta populasies geïsoleer is deur afstand. Die konklusie wat ek trek gebasseer op hierdie resultate is dat verskille in buis-lengtes in hierdie sisteem moontlik ontstaan het as gevolg van die verskillende bestuiwers wat aktief is in die verskillende T. revoluta populasies, en dat natuurlike prosesse nie die hoofrol spelers in hierdie sisteem is ten opsigte van buis-lengte evolusie nie.

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.