Thesis (MScConEcol (Conservation Ecology and Entomology))--University of Stellenbosch, 2011. / ENGLISH ABSTRACT: I evaluated the impact of Prosopis invasion and clearing on ecological structure, function and agricultural productivity in heavily grazed Nama Karoo rangeland on two sheep farms near the town of Beaufort West in the Western Cape Province of South Africa. My aims were to (1) determine the effects of invasion and clearing on rangeland vegetation composition, diversity (alien and indigenous species richness) and structure (alien and indigenous species cover), soil vegetation cover (plant canopy and basal cover) and agricultural productivity (grazing capacity), (2) describe the vegetation processes that underlay the invasion and clearing impacts and (3) evaluate the success of clearing in facilitating unaided restoration of ecological structure, function and agricultural productivity in formerly invaded rangeland. I hypothesised that invasion would significantly change rangeland vegetation composition and structure, leading to greater alien species richness and cover and lower indigenous species richness and cover while clearing would lead to lower alien species diversity and cover and greater indigenous species richness and cover. In addition I hypothesized that invasion would reduce rangeland plant canopy and basal cover and grazing capacity while clearing would substantially increase them. Finally I predicted that vegetation composition, alien and indigenous species cover and richness, plant canopy and basal cover and grazing capacity would revert to pre-invasion status and levels within four to six years of clearing. My results suggest that in heavily grazed Nama Karoo rangeland Prosopis invasion (~15 percent canopy cover) and clearing can significantly change rangeland vegetation composition, with invasion leading to greater alien species cover and lower indigenous species richness, while clearing leads to lower alien species richness and cover and greater indigenous species richness and cover. However invasion seems to have no effect on alien species richness and overall indigenous species cover. Clearing appears to facilitate the spontaneous restoration of alien species cover and indigenous species richness within four to six years but not species composition, alien species richness and indigenous species cover. In addition my results also indicate that Prosopis invasion can lower rangeland plant canopy and basal cover and grazing capacity while clearing, even under heavy grazing, can substantially raise them. Clearing however does not seem to facilitate the restoration of rangeland plant canopy and basal cover and grazing capacity to pre-invasion levels within four to six years after clearing. / AFRIKAANSE OPSOMMING: Ek het die impak van Prosopis indringing en verwydering van indringers op ekologiese struktuur, funksie en landbou produktiwiteit in ‘n swaar beweide Nama Karoo gebied op twee skaapplase naby Beaufort-Wes in die Wes-Kaap provinsie van Suid-Afrika geëvalueer. My doelwitte was om (1) te bepaal wat die gevolge van die indringing en verwydering van indringers op die natuurlike plantegroei samestelling, diversiteit (uitheemse en inheemse spesiesrykheid) en struktuur (uitheemse en inheemse spesies bedekking) sal wees, sowel as die effek op plantegroei bedekking (kroon en basalebedekking) en landbou produktiwiteit (weidingkapasiteit), (2) die plantegroei prosesse te beskryf wat onderliggend deur die impakte van indringing en verwydering van indringers veroorsaak word, en (3) die sukses van die verwydering van indringers te evalueer deur die fasilitering van blote restorasie van ekologiese struktuur en funksie en landbou produktiwiteit in voorheen ingedringde gebiede. My hipotese is dat indringing ‘n aansienlike verandering in natuurlike plantegroeisamestelling en struktuur sal veroorsaak, wat sal lei tot groter uitheemse spesiesrykheid en bedekking met minder inheemse spesiesrykheid en bedekking, terwyl die verwydering van indringers sou lei tot minder uitheemse spesie diversiteit en bedekking met 'n groter inheemse spesiesrykheid en bedekking. Verder vermoed ek dat indringing die natuurlike kroon- en basalebedekking en weidingkapasiteit sal verminder, terwyl die verwydering van indringers dit aansienlik sal verhoog. Ten slotte voorspel ek dat plantegroei samestelling, uitheemse en inheemse spesiesbedekking en -rykheid, kroon- en basalebedekking en weidingkapasiteit sou terugkeer na voor-indringing status en vlakke binne vier tot ses jaar na die verwydering van indringers. My resultate daarop dat die indringing van Prosopis (~ 15 persent kroonbedekking) en die verwydering van indringers in swaar beweide Nama Karoo gebiede ‘n aansienlike verandering in die gebied se natuurlike plantegroei samestelling toon, waar indringing gelei het tot groter uitheemse spesiesbedekking en minder inheemse spesiesrykheid, terwyl die verwydering van indringers lei tot minder uitheemse spesiesrykheid en groter inheemse spesiesrykheid en - bedekking. Dit lyk egter of indringing geen effek op uitheemse spesiesrykheid en algehele inheemse spesiesbedekking het nie. Die verwydering van indringers blyk om die spontane herstel van indringerbedekking en inheemse spesiesrykheid binne vier tot ses jaar te fasiliteer, maar nie spesiesamestelling, uitheemse spesiesrykheid of inheemse spesiesbedekking nie. Benewens dui my resultate ook aan dat Prosopis indringing die natuurlike kroon- en basalebedekking sowel as weidingskapasiteit verlaag, terwyl die verwydering van indringers, selfs onder swaar beweiding, die bedekking aansienlik kan verhoog. Verwydering van indringers lyk egter nie asof dit die herstel van die gebied se natuurlike kroon- en basalebedekking en weidingkapasiteit na voor-indringing vlakke toe kan fasiliteer binne vier tot ses jaar na die verwydering van indringers nie.

Mémoire numérisé par la Division de la gestion de documents et des archives de l'Université de Montréal.

Un modèle de croissance et de réponse à la radiothérapie pour le glioblastome multiforme (GBM) basé le formalisme du modèle de prolifération-invasion (PI) et du modèle linéaire-quadratique a été développé et implémenté. La géométrie spécifique au patient est considérée en modélisant, d'une part, les voies d'invasion possibles des GBM avec l'imagerie du tenseur de diffusion (DTI) et, d'autre part, les barrières à la propagation à partir des images anatomiques disponibles. La distribution de dose réelle reçue par un patient donné est appliquée telle quelle dans les simulations, en respectant l'horaire de traitement. Les paramètres libres du modèle (taux de prolifération, coefficient de diffusion, paramètres radiobiologiques) sont choisis aléatoirement à partir de distributions de valeurs plausibles. Un total de 400 ensembles de valeurs pour les paramètres libres sont ainsi choisis pour tous les patients, et une simulation de la croissance et de la réponse au traitement est effectuée pour chaque patient et chaque ensemble de paramètres. Un critère de récidive est appliqué sur les résultats de chaque simulation pour identifier un lieu probable de récidive (SPR). La superposition de tous les SPR obtenus pour un patient donné permet de définir la probabilité d'occurrence (OP). Il est démontré qu'il existe des valeurs de OP élevées pour tous les patients, impliquant que les résultats du modèle PI ne sont pas très sensibles aux valeurs des paramètres utilisés. Il est également démontré comment le formalisme développé dans cet ouvrage pourrait permettre de définir un volume cible personnalisé pour les traitements de radiothérapie du GBM. / We have developed and implemented a model of growth and response to radiotherapy for glioblastoma multiforme (GBM) based on the proliferation-invasion (PI) formalism and linear-quadratic model. We take into account patient-specific geometry to model the possible invasion pathways of GBM with diffusion tensor imaging (DTI) and the barriers to dispersal from anatomical images available. The actual dose distribution received by a given patient is applied as such in the simulation, respecting the treatment schedule. The free parameters in the model (proliferation rate, diffusion coefficient, radiobiological parameters) are randomly chosen from a distribution of plausible values. A total of 400 sets of values for the free parameters are thus chosen for all patients, and a simulation of the growth and the response to treatment is performed for each patient and each set of parameters. A failure criterion is applied to the results of each simulation to identify a site of potential recurrence (SPR). The superposition of all SPR obtained for a given patient defines the occurrence probability (OP). We show that high OP values exist for all patients and conclude that the PI model results are not very sensitive to the values of the parameters used. Finally, we show how the formalism developed in this work could help to define a custom target volume for radiation treatment of GBM.

L’Adénocarcinome Canalaire Pancréatique (ACP), la forme majeure de cancer du pancréas, est un des cancers les plus mortels du fait de son agressivité d’invasion locale et de dissémination vasculaire et de l’absence de méthode de détection précoce de la maladie. Nous avons développé un nouveau modèle d’invasion in vivo, sur la membrane chorio-allantoïdienne de l’embryon de poulet, permettant d’analyser les mécanismes d’interactions entre les cellules tumorales pancréatiques et leur microenvironnement. Nous avons montré que dans ce modèle les gènes codant pour les protéines sécrétées nétrine-1 et CXCL4L1/PF4v1 sont surexprimés dans les cellules tumorales au cours du processus d’invasion et que cette surexpression est également retrouvée dans les échantillons de patients humains. Nos études fonctionnelles ont indiqué que la nétrine-1 et CXCL4L1 pourrait jouer le rôle de régulateurs de la progression tumorale selon le modèle suivant : a) la chimiokine CXCL4L1 exercerait de façon paracrine une activité angiostatique sur les cellules endothéliales de l’ACP alors que b) la nétrine-1 aurait une activité pro-tumorale en agissant à la fois sur les cellules tumorales et sur les cellules endothéliales. Ces résultats ont permis d’une part de valider notre modèle en confirmant que les gènes surexprimés sélectionnés peuvent être impliqués dans la progression tumorale chez les patients. D’autre part, notre étude a permis de démontrer que les protéines CXCL4L1 et nétrine-1 constituent de nouvelles cibles thérapeutiques et/ou diagnostiques potentielles pour le cancer du pancréas. / Pancreatic Ductal Adenocarcinoma (PDAC), the major form of pancreatic cancer, is one of the deadliest cancers because of its propensity for local invasion and vascular dissemination and the lack of early diagnostic strategy. We have developed a new in vivo invasion model, on the chick embryo chorioallantoic membrane, allowing the analyze of mechanisms governing interactions between pancreatic tumor cells and their host microenvironment. We showed in its model that the genes encoding netrin-1 and CXCL4L1/PF4v1 secreted proteins are up-regulated in tumor cells in the course of the invasion process and we confirmed these up-regulation was also observed in human patients. Our functional studies indicated that netrin-1 and CXCL4L1 may play regulator roles in tumor progression according to the following model: a) CXCL4L1 chimiokine may have an angiostatic activity on endothelial cells by a paracrine mechanism of action whereas b) netrin-1 may have a pro-tumoral activity by acting on both endothelial and tumor cells. These results allowed in one hand to validate our model by showing that selected up-regulated genes may be involved in PDAC progression in human patients. On the other hand, our work provided evidence that CXCL4L1 and netrin-1 constitute new potential therapeutic and/or diagnostic targets for pancreatic cancer.

Die Leber ist das am häufigsten von Metastasen betroffene Organ und kann daher als Modellorgan für metastatische Invasion dienen. Aus diesem Grund war es das Ziel dieser Dissertation Genexpressionsprofile zu verstehen und metastasierungs- sowie invasionsassoziierte Gene zu identifizieren. Differentielle Genexpression wurde in drei Systemen überprüft: Einem syngenen Mausmodell, einem Xenograftmodell sowie in fünf Gewebeproben von Patienten. Genexpressionprofile des syngenen Mausmodells und der Patientenproben zeigten, dass man die Invasionsfront als Ganzes betrachten, um möglichst viele über-lappende Gene zu finden. Globale Genexpressionstudien, die auf den Wirtsteil der Invasionsfront zeigten bemerkenswerte Überrepräsentation z. B. der „GO-terms“ „extrazelluläre Matrix“, Zellkommunikation“, „Antwort auf biotischen Stimulus“, Strukturmolekülaktivität“ und „Zellwachstum“. Marker der Aktivierung hepatischer Sternzellen überrepräsentiert in der invasionsfront, was die Durchführbarkeit einer Analyse differentieller Genexpression im genomweiten Rahmen anzeigt. Globale Genexpressionsstudien, auf den Tumorzellen in der in vitro Situation, in vivo und in der Invasionsfront zeigten insgesamt einen Anstieg zellulärer Spezialisierung von der in vitro zur Invasionsfront. Sezernierte proangiogenetische Chemokine zeigten eine Hochregulation in der Invasionsfront. Das beta catenin Gen war in der Invasionsfront 9.6 fach erhöht im Vergleich zur in vitro Situation. Die Überprüfung der transkriptionellen Aktivierung von beta catenin über die Prüfung der Promotoraktivität zeigte einen 18.4 fachen Anstieg in den Tumorzellen der Invasionsfront. Weiterhin war die Promotoraktivität (an Hand der Aktivität der mRNA des Alkalischen Phosphatase Reportergens) im Tumorinneren 3.5 fach höher als in der Zellkultur, was für einen transkriptionellen Mechanismus der beta catenin Regulation zusätzlich zu den posttranslationalen Mechanismen spricht. / Liver is most frequently populated by metastases and may therefore serve as a model organ for metastatic invasion. So the aim of this thesis is to understand the gene expression profiles and identify metastasis and invasion related genes. Differential gene expression was examined in three systems: A syngeneic mouse model, a xenograft model and five clinical specimens. Gene expression profiles of a syngenic mouse model and human clinical specimen revealed that the invasion front should be considered as a whole to find more overlapping potential target genes. Global gene expression studies on the host part of the invasion front, revealed a pronounced overrepresentation of GO-terms (e.g. “extracellular matrix”, “cell communication”, “response to biotic stimulus”, “structural molecule activity” and “cell growth”). Hepatic stellate cell activation markers were over-represented in the invasion front demonstrating the feasibility of a differential gene expression approach on a genome wide scale. Global gene expression studies of the tumor cells in vitro, in vivo and tumor part of the invasion front revealed an overall increase of cellular specialization from in vitro to the invasion front. Secreted angiogenic cytokines were found to be up regulated in the invasion front. Beta catenin gene of “cell adhesion” GO term was elevated 9.6 fold in invasion front compared to in vitro. Evaluation of transcriptional up-regulation of beta catenin by promoter activity showed an 18.4 fold increase in the tumor cells of the invasion front as compared to those from the faraway tumor. Promoter activity assessed by soluble human placental alkaline phosphatase reporter gene mRNA was 3.5 fold higher in the inner parts of the tumor than in vitro cells indicating a transcriptional mechanism of beta catenin regulation in addition to the posttranslational regulatory mechanisms.

L’abeille mellifère (Apis mellifera) est une espèce clé dans son aire d’origine tant pour les services écosystémiques rendus que pour les productions apicoles. Toutefois, ses populations sont soumises à différents facteurs de stress biotiques responsables de mortalités importantes. Dans les îles du Sud- Ouest de l’Océan Indien (SOOI) la sous-espèce d’abeille endémique est A. m. unicolor. Toutefois, aucune étude n’avait été réalisée sur sa pathosphère et ce n’est que depuis 2010 où l’ectoparasite Varroa destructor a envahi certaines de ces îles, menaçant cette sous-espèce, que de premières études ont été conduites. Dans ce travail de thèse, nous avons réalisé un état des lieux général de la santé des cheptels d’abeilles grâce à des enquêtes épidémiologiques descriptives dans la majorité des îles : La Réunion, Maurice, Rodrigues, Madagascar et les Seychelles. Nous avons pu montrer des faciès épidémiologiques assez similaires entre les îles, marqués par une dominance de Nosema ceranae particulièrement dans les petites îles (80-100%) et la présence de 3 virus : BQCV (4-89%), CBPV (2-51% excepté à Rodrigues) et DWV (4-40% excepté à La Réunion). D’autres agents pathogènes n’ont été retrouvés que sur certains territoires comme Aethina tumida, Braula pretoriensis, Acarapis sp. ou Melissococcus plutonius. L’analyse de la diversité génétique réalisée sur les 3 virus a montré une proximité des souches virales au sein du SOOI. Les enquêtes réalisées dans un contexte sans varroa ont montré une bonne santé des colonies avec une absence de signes cliniques. Les mortalités constatées n’ont concernées que les territoires envahis par varroa : Madagascar, Maurice, La Réunion. Varroa semble donc être le principal responsable des mortalités aiguës de colonies dans la zone bien avant les autres agents pathogènes ou les causes environnementales. Son impact sur les populations d’abeilles et in fine sur les écosystèmes indigènes sera à évaluer dans les années futures. / The honeybee (Apis mellifera) is a key species in its native range for both ecosystem services offerred and for bee products. However, its populations are subject to various biotic stressors responsible for significant mortalities. In the South-West Indian Ocean region the endemic bee subspecies is A. m.unicolor. However, no studies had been carried out on its pathosphere and it is only since 2010 where the ectoparasite Varroa destructor invaded some of these islands, threatening this subspecies, that first studies were conducted. In this thesis work, we carried out a general inventory of the health of honeybee herds through descriptive epidemiological surveys in the majority of islands: Réunion, Mauritius, Rodrigues, Madagascar and Seychelles. We were able to show quite similar epidemiological facies between islands, characterized by a dominance of Nosema ceranae especially in small islands (80-100%) and the presence of 3 viruses: BQCV (4-89%), CBPV (2- 51% except in Rodrigues) and DWV (4-40% except in Reunion). Other pathogens have only been found in certain territories such as Aethinatumida, Braula pretoriensis, Acarapis sp. or Melissococcus plutonius. The analysis of the genetic diversity carried out on the 3 viruses showed a proximity of viral strains within the SOOI. Surveys conducted in a context without varroa showed good colony health with no clinical signs. The observed mortalities concerned only the territories invaded by varroa: Madagascar, Mauritius, Reunion. Varroa therefore appears to be the main cause of acute colony mortality in the area long before other pathogens or environmental causes. Its impact on bee populations and ultimately on native ecosystems will be evaluated in future years.

There are many factors that have been proposed to contribute to plant invasiveness in nonnative ecosystems. Traits of invading species are one of them. It has been proposed that successful species at a certain invasion stage share particular traits, which could be used to predict the behaviour of potentially invasive plants at the respective stage. Three main stages of invasion are distinguished: introduction, naturalization, and invasion. I conducted a stageand trait-based analysis of available data for the invasion of New Zealand by the flora of southern Africa. Using 3076 southern African native vascular plant species introduced into New Zealand, generalised linear mixed model analysis was conducted to assess association of several species traits with the three invasion stages. The results showed that plant traits were significantly associated with introduction but fewer traits were associated with naturalization or invasion, suggesting that introduction can be predicted better using plant traits. It has been also hypothesized that species may become invasive in non-native ecosystems because they are removed from the regulatory effects of coevolved natural enemies (Enemy Release hypothesis). A detailed field study of the succulent plant Cotyledon orbiculata var. orbiculata L. (Crassulaceae) was conducted in the non-native New Zealand and native Namibian habitats to compare the extent of damage by herbivores and pathogens. C. orbiculata is a southern African species that is currently thriving in New Zealand in areas seemingly beyond the climatic conditions in its native range (occurring in higher rainfall areas in New Zealand than are represented in its native range). As hypothesised, C. orbiculata was less damaged by herbivores in New Zealand but, contrary to expectation, more infected by pathogens. Consequently, the plant was overall not any less damaged by natural enemies in the non-native habitat than in its native habitat, although the fitness impacts of the enemy damage in the native and invaded ranges were not assessed. The results suggest that climatic conditions may counteract enemy release, especially in situations where pathogens are more prevalent in areas of higher rainfall and humidity. To forecast plant invasions, it is concluded that species traits offer some potential, particularly at the early stage of invasion. Predicting which introduced plants will become weeds is more difficult. Enemy release may explain some invasions, but climatic factors may offset the predictability of release from natural enemies.

Under senare tid har allt fler företag börjat utöva verksamhet där mark nyttjas för olika aktiviteter med stöd av allemansrätten. HD har bedömt att sådant nyttjande; kanalisering, är tillåtet med stöd av allemansrätten, men uttalade i NJA 1996 s. 495 (”Forsränningsmålet”) samtidigt att nyttjandet bara är tillåtet så länge marken inte skadas. Eftersom skadeståndsfrågan aldrig har prövats i samband med allemansrätten har föremålet för denna uppsats varit att utreda möjligheterna att utfå ersättning för skador via allmänna skadeståndsrättsliga regler, skadestånds- och avhjälpandeansvaret i miljöbalken samt för intrång i äganderätten via expropriationslagen och möjligheten att få ersättning ur en försäkring. Expropriationsalternativet kan inte användas idag, eftersom marknyttjande med stöd av allemansrätten är ett icke ersättningsgillt intrång i äganderätten. Möjligheterna till att betrakta en skada på marken som en sakskada är stora. Det finns emellertid omständigheter som skapar komplikationer för båda skadeståndsansvarsformerna. En allmän skadeståndstalan kräver att culpa hos skadevållaren kan bevisas, vilket kan vara svårt när många människor vistas på en plats samtidigt. Kanalisatören kan betraktas som culpös enligt principerna om kumulativ eller anonym culpa, enligt vilka en principal blir ansvarig, trots att de individer han ansvarar för inte, var och en för sig, varit culpösa. Det miljörättsliga skadeståndsansvaret bygger på ersättning för störningar som inte är orts- eller allmänvanliga. När det gäller allemansrättsligt nyttjande torde många störningar betraktas som just orts- eller allmänvanliga. Avhjälpandeansvaret i miljöbalken bygger på ersättning för miljöfarlig verksamhet, och det är tveksamt om det kan tillämpas på kanalisatörens verksamhet. Expropriationsersättning på grund av allemansrättsligt nyttjande tillerkänns inte markägaren enligt gällande rätt, vilket kan tyckas märkligt eftersom många nyttjandeformer som påminner om allemansrätten, t.ex. servitut, medför sådan ersättningsrätt. Det finns vissa möjligheter att ersätta markskador ur försäkringar. Dock är de flesta skador följden av invasionsproblematik, som är mycket svårt försäkra. Samtliga ersättningsformer har visat sig otillräckliga för att ersätta skador som är estetiska och ideella, eftersom sådana skador inte kan beräknas i pengar. Beroende på hur marknyttjande med stöd av allemansrätten betraktas har man olika åsikt om i vad mån kanalisatören borde ersätta markägaren för skador på marken. Om nyttjandet uppfattas som någonting som finns vid sidan av markäganderätten är endast konkreta skador ersättningsgilla. Om man anser att nyttjandet är ett intrång i äganderätten och en förmögenhetsöverföring från markägaren till kanalisatören bör emellertid redan rätten att nyttja marken föranleda ersättning. Den ersättningsform som bäst tillgodoser markägarens intresse av att få skador på marken åtgärdade och få ersättning för ekonomisk förlust är avhjälpandeansvaret. Endast det ansvaret reparerar till fullo skador på miljön. Det krävs emellertid ett domstolsavgörande för att slutligen klargöra avhjälpandeansvarets tillämplighet. / Recently, more and more companies have come to exercise activities under the right of public access on land that belongs to whomever. The High Court has assessed that such use of land; channeling, is permitted, but in NJA 1996 s. 495 (“The Rafting Case”) also announced that this use is only allowed as long as the land remains unharmed. Since the question of damages in the context of the right of public access has not yet been treated in jurisprudence, the object of this thesis has been to investigate the possibilities to obtain restitution for damages via tort law, liability under the environment act as well as for proprietary intrusions under the expropriation act and restitution via insurance. Expropriation, however, cannot currently be made use of, since the right of public access is an intrusion which does not entail restitution. There are extensive possibilities for considering damage to the land as damage to property. However, there are ramifications for both the legal options for damages: the tort law and the environmental act. The tort law requires tortuous behaviour from the tortfeasor to be established, which is difficult when several persons stay at one place simultaneously. The canalizer may be considered tortuous under the principle of vicarious liability and cumulative or anonymous tort. According to these principles, the canalizer is responsible for the actions of certain individuals, even though they have not, individually, been negligent. The environmental liability for damage is based on restitution for disturbances that are not common to the place nor to the general public. This fact is difficult to establish in the context of the right of public access. The remedial responsibility of the environmental act is constructed for operations that are harmful to the environment, which is why it is dubious whether the responsibility is applicable to the canalizer’s activities. Currently, compensation for expropriation cannot result of use of land under the right of public access. This is peculiar, since many usufructs reminiscent of the right of public access, i.e. the easement, result in such compensation. Some forms of damage to the land can be insured. Although, most damage in this context arise from invasion, a cause of damage which is nearly impossible to insure. All forms of restitution have proven insufficient for compensating damage that is esthetic or otherwise non-pecuniary. Depending on if the right of public access is regarded as proprietary intrusion or something that exists alongside the right to property, one comes to different conclusions about the extent of compensation for damage. In the former case, only substantial damage is compensable. In the latter case, even the right to use the land occasions compensation. The liability which best meets the needs of the proprietor: compensation and restoration of damage to his property, is the remedial responsibility of the environmental code. This responsibility alone fully restores harm to the environment. However, the applicability of the remedial responsibility needs to be established in jurisprudence.

The spread of exotic species is considered to be one of the most significant threats to ecosystems and emphasises the need for appropriate management interventions. The majority of marine non-indigenous species (NIS) are believed to have been introduced via ship biofouling and their domestic spread continues to take place via this mechanism. In some countries, biosecurity systems have been developed to prevent the introduction of NIS through biofouling. However, implementing biosecurity strategies is difficult due to the challenges around identifying high-risk vectors. Reliable predictors of risk have remained elusive, in part due to a lack of scientific knowledge. Nonetheless, invasion ecology is an active scientific field that aims to build this knowledge. Propagule pressure is of particular interest in invasion ecology as it describes the quantity and quality of the propagules introduced into a recipient region and is considered to be an important determinant in the successful establishment of NIS. Environmental history affects health and reproductive output of an organism and, therefore, it is beneficial to examine this experimentally in the context of biofouling and propagule pressure. The aim of this thesis was to examine how voyage characteristics influence biofouling recruitment, survivorship, growth, reproduction and offspring performance through the ship invasion pathway. This was to provide fundamental knowledge to assist managers with identifying high-risk vessels that are likely to facilitate the introduction or domestic spread of NIS, and to understand the processes affecting biofouling organisms during long-distance dispersal events. Chapter One provides an introduction to the issues addressed in this thesis. Each data chapter (Chapters Two – Five) then focused on a stage of the invasion process and included field experiments using a model organism, Bugula neritina. Finally, Chapter Six provides a summary of key findings, discussion and the implications to biosecurity management. Throughout this thesis, the effect of donor port residency period on the success of recruits was highlighted. Chapter Two focused on recruitment in the donor region. As expected, recruitment increased with residency period. Importantly, recruitment occurred every day on vulnerable surfaces, therefore, periods as short as only a few days are able to entrain recruits to a vessel hull. The study presented in Chapter Three showed that there was high survivorship of B. neritina recruits during 12 translocation scenarios tested. In particular, the juvenile short-residency recruits (1-8 days) survived voyages of 8 days at a speed of 18 knots; the longest and fastest voyage simulated. Interestingly, variation in voyage speed and voyage duration had no effect on the survivorship of recruits, but did have legacy effects on post-voyage growth. Again, B. neritina which recruited over very short residency periods of 1 day continued to perform well after translocation and had the highest level of reproductive output after the voyage scenarios (Chapter Four). Recruits that were older (32-days) and reproductively mature at the commencement of the scenarios failed to release any propagules. Even though the number of ‘at sea’ and ‘port residency’ days were equal, reproductive output was higher after short and frequent voyages than after long and infrequent voyages. Finally, the study presented in Chapter Five examined transgenerational effects of B. nertina. Results showed that although the environmental history of the parent colony had a carry-over effect on offspring performance, it was the offspring environment that was a stronger determinant of success (measured by reproductive output and growth). Although cross-vector spread is possible (i.e. parent and offspring both fouling an active vessel), offspring released from a hull fouling parent into a recipient environment will perform better. In combination, these studies have provided new insights into NIS transport via vessel biofouling. Although shipping pathways are dynamic and complex, these results suggest that juvenile stages that recruit over short residency periods and are then translocated on short voyages, may pose a higher risk for NIS introduction than originally assumed. This has implications for marine biosecurity management as short residency periods are common and short, frequent voyages are typical of domestic vessel movements which are largely unmanaged.

No description available.