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.

MSc., Faculty of Science, University of the Witwatersrand, 2011 / Both indigenous “Tamarix usneoides” and alien T. ramosissima co-occur in South Africa. Tamarix usneoides is potentially a metal and salt phytoremediation species. However, hybridisation is occurring between the two species and hybrids growth characteristics are deemed to be invasive. This study was undertaken at two sites on one Highveld gold mine, characterised by different soil types, where plant growth, reproductive potential and insect abundance and diversity were measured. This aims to establish the growth of T. usneoides in two site-species trials, and to identify differences between Tamarix taxa that are consistent across different environmental conditions and can be used for identification. This study also set out to establish if higher reproductive effort and enemy release are promoting the invasiveness of T. ramosissima. Trees in one site (a water logged, clayey “Glenrosa” soil) displayed significantly greater percentage survival, growth and reproductive effort than trees in the other site (a well drained, sandy “Hutton” soil). Extensive hybridisation between T. usneoides and T. ramosissima was observed with two hybrid states, morphologically similar to each parent species. The reproductive effort is consistently greater in T. ramosissima and T. ramosissima hybrid than T. usneoides and T. usneoides hybrid. After sampling 217 trees in this study, the three characteristics of (1) insertion of filaments into the nectar disc, (2) petal shape and (3) presence of salt glands on leaves most accurately distinguished between T. usneoides and T. ramosissima taxa. Three insect species have been identified from this study as potentially damaging agents on Tamarix spp. Insects can discern between the Tamarix tree taxa and T. ramosissima is being released from insect herbivore pressure, as very few insects and a lower diversity were recorded on T. ramosissima and its hybrids (T. ramosissima hybrids > T. ramosissima). To achieve phytoremediation objectives, careful tree identification and selection of genetically pure individuals for cloning is recommended. Additionally the removal of existing alien and hybrid plants, and further research to find a suitable host specific insect biological agent to control alien Tamarix is necessary.

O Cerrado é o bioma brasileiro que mais perdeu área para o agronegócio em anos recentes, verificando que a expansão da fronteira agrícola sobre o Cerrado provocou a pulverização dos remanescentes em fragmentos cada vez menores, imersos em matrizes de baixa similaridade, causando uma série de impactos negativos sobre o ecossistema natural. A fragmentação de habitats é considerada uma das mais graves ameaças à conservação da biodiversidade, com consequências ecológicas variáveis, intensificadas na faixa de transição entre matriz e fragmento. Os efeitos de borda são decorrentes de mudanças físicas e bióticas nas faixas marginais das ilhas de vegetação natural, que promovem modificações nas características microclimáticas, as quais conduzem a uma série de alterações importantes nas comunidades que compõem os ecossistemas. O objetivo deste estudo foi investigar a existência de efeito de borda e suas consequências sobre a comunidade vegetal do Cerrado stricto sensu. O estudo foi realizado em fragmento de Cerrado com área de 980,8 ha, localizado no município de Iaras, SP, cujo entorno foi ocupado por pastagem durante cerca de um século. Efetuou-se a caracterização estrutural e florística de diferentes estratos da vegetação em diferentes distâncias da borda (0, 10, 20, 40, 80 m) e coletaram-se dados microclimáticos na matriz (a 10 m de distância da borda) e no interior do fragmento, nas diferentes distâncias da borda e em duas estações do ano (seca e chuvosa). Investigou-se a existência de um possível gradiente de estrutura ou composição da vegetação relacionado com a distância da borda e com as variáveis microclimáticas. Os resultados obtidos mostraram não existir variações de microclima e de estrutura e composição do componente arbóreo e herbáceo-arbustivo em relação à distância da borda do fragmento. Embora independente do microclima e, portanto, diferente dos clássicos efeitos de borda, a invasão pela braquiária (Urochloa decumbens) foi detectada como a única consequência da exposição da periferia do Cerrado stricto sensu às pressões da matriz, causando exclusão das gramíneas nativas e inibindo o desenvolvimento de plantas lenhosas de pequeno porte. A ausência de efeito de borda sobre os demais componentes da vegetação sugere, em um primeiro momento, que esta não é uma ameaça importante para a conservação do cerrado. Porém, a invasão biológica pela gramínea africana é relativamente recente (foi introduzida há cerca de três décadas na propriedade) e suas consequências ainda não podem ser avaliadas adequadamente. É possível que seus impactos sobre os estratos superiores da vegetação venham a ser observados no futuro, quando a inibição da regeneração das espécies arbustivas e arbóreas comprometer a dinâmica da comunidade vegetal e a substituição dos adultos nas populações dessas espécies. Além disso, hoje a invasão só causa impactos significativos sobre a faixa de 0 a 20 m de distância da borda, mas não é possível prever se a gramínea invasora vai se expandir rumo ao núcleo do fragmento no futuro. / From the Brazilian biomes, the Cerrado has been the most widely and rapidly converted to agribusiness in recent years. The expansion of agricultural frontier over the Cerrado domain resulted in thousands of small fragments embedded in matrices of low similarity, causing a series of negative impacts on the natural ecosystem. Habitat fragmentation is considered one of the most serious threats to the conservation of biodiversity and ecological consequences vary, being intensified in the range of transition between matrix and fragment. The edge effects are caused by biotic and physical changes in marginal areas of the islands of natural vegetation, promoting changes in microclimatic characteristics, which leads to a series of important changes in the ecosystems. The aim of this study was to investigate the existence of edge effect and their consequences on the plant community of Cerrado sensu stricto. The study was conducted in Cerrado fragment with an area of 980.8 ha, located in the municipality of Iaras, SP, whose surrounding area was occupied by pasture for about a century. Structure and floristic composition of distinct vegetation layers was assessed at different distances from the edge (0, 10, 20, 40 and 80 m). Climatic data were collected in two seasons (dry and wet), at the same distances withing the fragment and also in the matrix (10 m away from the edge). The existence of a gradient of vegetation composition or structure and the microclimatic variables related to distance from the edge was investigated. No changes in microclimate and structure or composition of the wood layers related to distance from the edge was observed. Although independent of the microclimate and therefore different from classic edge effects, invasion by the African grass Urochloa decumbens (braquiária) was detected as the only consequence of exposure of the periphery of the cerrado sensu stricto to the pressures of the matrix, causing the exclusion of native grasses and inhibiting the development of small woody plants. The absence of edge effect on other components of the vegetation suggests, at first, that this is not a major threat to the conservation of the Cerrado. However, the biological invasion by the African grass is relatively recent (it was locally introduced about three decades ago) and its consequences have not been adequately assessed. It is possible that their impacts on the upper strata of the vegetation may be observed in future, when the inhibition of regeneration of shrub and tree species jeopardize the dynamics of plant community and the replacement of adults in the populations of these species. In addition, at present the invasion has significant impacts only over 0-20 m from the edge, but it is impossible to predict whether the invasive grass will expand or not into the core area of the fragment in the future.

The British Isles have been invaded by 12 alien cynipid gallwasps over the past 150 years. The first 4 of these species have been studied in depth and represent a model system in phytophagous insect community structure. In this thesis, I extend this research programme to incorporate 8 further invaders. I examine recent changes in the distribution of invading oak gallwasps in Britain and spatial patterns in the composition of the associated communities of phytophagous cynipid inquilines and parasitoids. I use fully quantitative webs to assess the diversity and strength of trophic interactions between native and invading species and assess the potential for apparent competition between gallwasps mediated by shared natural enemies. Of the first 4 invaders to be studied, 3 have expanded their range since 1991/2. Three of these 4 species are now well established in Scotland, while Andricus corruptrix remains confined to England. Four new invaders (A. aries, A. lucidus, A. grossulariae, Aphelonyx cerricola) are established in southern England and are spreading. Rates of range expansion vary across species (between means of 3.3 and 24.4 km per year), and may be correlated with variation in lifecycles and abundance. The four newest invaders (Neuroterus saliens, Plagiotrochus australis, P. coriaceus, P. quercusilicis) are currently restricted to their sites of first record. Previous studies on one of the early invaders, Andricus quercuscalicis, identified south to north and east to west declines in community species richness and in the abundance of specific parasitoid species. I find that: 1) Parasitoid associations with the asexual galls of A. quercuscalicis track inquiline recruitment to this host. 2) The longitudinal and latitudinal gradients in parasitoid species richness demonstrated in previous work are no longer apparent, suggesting that younger northern communities may be converging on their older southern counterparts. 3) Inquilines show increasing survivorship with distance from the original centre of their distribution in south east England, suggesting at least temporary exploitation of enemy-free space. 4) The recently invading Andricus and Aphelonyx species have all rapidly recruited parasitoids and inquilines. Fully quantitative webs were constructed for 4 sites in England and Scotland incorporating both native and invading cynipids. I tested the hypothesis that newly arriving gallwasp generations would fall within food web compartments based on their host oaks and location on the tree as demonstrated in previous work. Counter to this hypothesis, parasitoids attacking one of the newest invaders (A. grossulariae) break down host tree-associated compartmentalisation. Where A. grossulariae has yet to become established, host-based compartmentalisation remains pronounced. Despite extensive sharing of parasitoid species, I found only one strong indirect interaction between species (both aliens) and no evidence for widespread apparent competition. Spatial density dependent predation on an appropriate scale can stabilise population dynamics. I quantified predation by blue tits (Parus caeruleus) of spring generation bud galls on Turkey oak (Q. cerris) at three spatial scales (shoots within branches, branches within trees, trees within sites). I found significant levels of bird predation, with most variation occurring between trees rather than between shoots within branches. Spatial density dependence was detected at sites in southern England, primarily at the level of trees within a site. Relationships at finer spatial scales were far more variable in magnitude and sign. My results suggest that blue tits forage primarily at the level of trees. This thesis presents comprehensive new data on the establishment and spread of 12 invading cynipid species and on their interactions with native communities. The results further understanding of both spatial and temporal aspects of natural enemy recruitment to invading species. In particular, it is clear that individual invading species can significantly modify trophic linkage between established food web compartments. Finally, my data emphasise the significant (but often unstudied) contribution of highly mobile vertebrate predators to otherwise closed ecological microcosms.

Thesis (doctoral)--Swedish University of Agricultural Sciences, 2007. / Thesis documentation sheet inserted. Appendix reproduces five papers and manuscripts, two co-authored with others. Includes bibliographical references. Also issued electronically via World Wide Web in PDF format; online version lacks appendix.

Plant invasions can ensue when plants are introduced to regions without their specialist enemies (the Enemy Release Hypothesis). This assumes natural enemies limit survival and fecundity in an invader's native range. I tested this assumption for a native invasive species, Common Ragweed (Ambrosia artemisiifolia), by excluding natural enemies from seeds, seedlings and adults. At the adult stage, I added disturbance and conspecific density treatments. Protection from herbivores slightly improved performance at the seedling stage only, while disturbance greatly increased survival and fecundity. Increasing conspecific density reduced performance only in disturbed plots. I also tested herbivore tolerance using simulated damage. Heavy (75%) damage did not reduce fecundity; light damage even increased seed production. These results suggest enemies do not limit Ambrosia in its native range, especially compared to the effects of habitat disturbance. While enemy release may have occurred during Ambrosia’s invasions, it is not likely to be their principal cause.

Plant invasions can ensue when plants are introduced to regions without their specialist enemies (the Enemy Release Hypothesis). This assumes natural enemies limit survival and fecundity in an invader's native range. I tested this assumption for a native invasive species, Common Ragweed (Ambrosia artemisiifolia), by excluding natural enemies from seeds, seedlings and adults. At the adult stage, I added disturbance and conspecific density treatments. Protection from herbivores slightly improved performance at the seedling stage only, while disturbance greatly increased survival and fecundity. Increasing conspecific density reduced performance only in disturbed plots. I also tested herbivore tolerance using simulated damage. Heavy (75%) damage did not reduce fecundity; light damage even increased seed production. These results suggest enemies do not limit Ambrosia in its native range, especially compared to the effects of habitat disturbance. While enemy release may have occurred during Ambrosia’s invasions, it is not likely to be their principal cause.

Aquest treball de tesis doctoral tracta sobre la formiga invasora Lasius neglectus i els seus efectes sobre altres artròpodes en comunitats envaïdes al Nord-est de la Península Ibèrica. S'han estudiat: a) trets de la seva biologia reproductiva i ecologia b) els efectes que la invasió provoca sobre altres espècies de formigues natives i sobre els altres artròpodes que coexisteixen amb la colònia de L. neglectus; aquest darrer aspecte s'ha estudiat en tres poblacions. També es va fer un assaig de control de la plaga amb un tractament clàssic amb insecticides de contacte i una elevada persistència. Pels estudis sobre la seva biologia reproductiva es va constatar el seu caràcter poligínic en tota la supercòlonia, amb una gran densitat de reines per supercolònia que es trobaven isolades o bé en grups de 2 o més reines dins de cada niu i sempre amb obreres. Per les dades en increment en pes sec i contingut de greixos des del naixement fins al moment de l'aparellament es va demostrar que les reines de L. neglectus es poden reproduir mitjançant fundació dependent tot i que el valor obtingut de la proporció del mesosoma de reines i obreres (relació entre el volum del tòrax de reines i obreres=23.83±0.15) coincideix amb el de les espècies que fan fundacions independents. Al laboratori també es van obtenir obreres a partir de reines joves acabades de fecundar (fundacions independents, haplo i pleometrótiques). Per les dades en contingut de carbohidrats es demostra que les reines tenen valors entremitjos entre les reines d'espècies que fan vols nupcials i les que no són capaces de volar, tot i que per l'àrea relativa de les ales les reines de L. neglectus han de ser capaces de volar. Tanmateix mai s'ha documentat cap vol nupcial en L. neglectus fins ara. Les reines joves alades fan fecundació intranidal i aquestes reines joves fecundades són adoptades per les reines velles i les obreres dins del mateix niu. Aquesta estratègia evita la depredació de reines joves fecundades després del vol nupcial, moment crític per a l'expansió de la colònia o la dispersió de l'espècie. Per tant aquesta expansió és per gemmació o bé a través de transport passiu, normalment afavorit per l'home. Les taxes de posta de reines velles són el doble que les de reines joves, amb un temps mitjà de desenvolupament de les obreres, però, el doble de llarg. L'efecte més significatiu de la invasió de L. neglectus, tant a nivell d'abundància com de riquesa d'espècies, es va trobar sobre la resta de formigues natives. En totes les poblacions estudiades la presència de L. neglectus reduïa significativament la probabilitat de trobar espècies de formigues natives tant al sòl com als arbres. En el cas de l'efecte sobre altres artròpodes els resultats eren més variables: alguns artròpodes com ara els àfids es veien afavorits per la seva presència i en canvi altres ordres com ara Aràcnids, Himenòpters i Dípters es veien afectats negativament per la presència de la formiga invasora. Tanmateix aquests efectes, tant en formigues natives com en altres artròpodes, variaven en funció de la població estudiada i depenent de l'hàbitat mostrejat: capçades dels arbres o bé sòl.En quan a les mesures per controlar aquesta invasió, es va fer un assaig amb insecticides d'una colònia de L. neglectus. Es van atacar tots els fronts possibles d'expansió de la invasió: a) l'interior de cases ocupades amb esquers, b) el perímetre de les cases ocupades amb injeccions amb un insecticida de contacte amb gran persistència al igual que c) els troncs dels arbres i finalment, d) es van tractar totes les capçades dels arbres de la zona ocupada per tal de matar els àfids, que són el principal aliment de L. neglectus. Gràcies al coneixement de les dades biològiques i ecològiques de la supercolònia de L. neglectus el resultat d'aquests tractaments integrats va ser satisfactori ja que disminuïa dràsticament la probabilitat de trobar formigues després del tractament i, a l'any següent al tractament, els nivells inicials de formigues eren molt menors als de l'any abans de tractar. Els tractaments es van fer dos anys seguits amb resultats prometedors i gran satisfacció per part dels habitants de les cases ocupades. / The aim of this work is to study: a) the reproductive biology and ecology of the invasive garden ant Lasius neglectus and b) its effects over other native arthropod communities in invaded communities from Northeast Spain; this last aspect has been studied on three populations. A large scale insecticide treatment of the plague was also made with contact and high persistence insecticides. Its polygynic character was ascertained by the studies about its reproductive biology, with a great density of queens that were found isolated or in groups of 2 or more queens per nest, always with workers. From data obtained in increase in dry weight and fat contents from eclosion until mating time, it was demonstrated that L. neglectus queens can reproduce through dependent colony founding even though the value obtained from the proportion of the mesosome ratio from queens and workers (queen/worker thorax volume=23.83±0.15) coincides with the independent colony founding species. Workers were also obtained, at the laboratory, from newly mated queens (haplo- and pleometrotic independent foundations). From data in carbohydrate content, queens have intermediate values among the flying queens and those not capable to perform nuptial flights, even though for the relative area of the wings, L. neglectus queens are capable of flying. However it has never been documented a nuptial flight in L. neglectus until now. Queens reproduce by intranidal mating and these fertilized young queens are adopted by the old queens and workers into the same nest. This strategy avoids the possible predation of young fertilized queens after the nuptial flight, a critical moment for the expansion of the colony. Therefore they expand by budding or through passive human transport. The egg laying rates of old queens are two fold than that of young mated queens, with a total development time from egg to worker double than from egg to nanitic for young queens. The most significant effect of the invasion of L. neglectus, on native arthropods abundance and species richness, was on coexisting native ants. In all the studied populations the presence of L. neglectus reduced dramatically the likelihood of finding native ants on tree canopies and soil. For the effect of L. neglectus on other non-ant arthropods the results were more variable: some arthropods like aphids were favoured by its presence and other orders as Arachnids, Hymenoptera and Diptera were negatively affected for the presence of the invading ant. However these effects, in native ants as well as in other arthropods, vary depending on the studied population and on the sampled habitat: tree canopy or soil surface. In order to control this invasion, classical treatments with insecticides were applied to one of the supercolonies studied. All the possible expansion fronts of the invasion were attacked with a) an in-house baiting, b) a house perimeter injection treatment, c) tree trunk spraying, and finally d) canopy fogging in order to kill the aphids, which are the main food supply for L. neglectus. Thanks to the knowledge of the biological and ecological features of the supercolony of L. neglectus the results of these integrated treatments were satisfactory. They drastically reduced the probability of finding ants after the treatment and, for the next year after the first treatment, the initial ant levels were lower than those of the year before treating. This treatment was repeated for two years with promising results and great satisfaction of the homeowners.

This project analyzes two sets of alien invasion films to understand lay opinions on hegemonic authority. It defines hegemonic authority along two major lines: neo-Gramscian hegemony and hegemonic stability theory. The project uses alien invasion films to study challenges to hegemonic authority because of the unique and confrontational narrative alien invasion films typically possess. Through a comparative process, the project concludes that alien invasion films reveal paradoxical relationships of power, where the hegemon encourages aggressive pre-emptive policies against its challengers but at the same time depends upon these challengers to maintain its power. It argues that despite arguments of growing globalism and cosmopolitanism in the world, the liberal hegemon remains clearly divided among notions of Us versus Them.

Thesis (Ph. D.)--University of California, San Diego, 1998. / Includes bibliographical references (p. 197-209).