Many regional and local factors can influence the distribution of native and exotic species in ecological communities. I examined the regional- and local-scale determinants of native and exotic vascular plant species richness and composition in a highly fragmented oak savanna ecosystem on southeastern Vancouver Island. In sharp contrast to most reported results, I found a negative relationship between native and exotic richness at the regional scale, and no relationship at the local scale. Two extrinsic factors, surrounding road density and climate, best explained the regional-scale relationship by each affecting natives and exotics in opposite ways. Road density and climate were also the dominant predictors of native and exotic composition at the regional scale. Patterns in the patch occupancy of individual species confirmed the importance of these factors but I found that low surrounding road densities and cool, wet conditions predicted the presence of many natives and the absence of many exotics. Environmental factors explained variation in richness and composition at the local scale, but these factors were different for natives and exotics. My results suggest that natives and exotics respond to roads and climate in fundamentally different ways. Roads increase both exotic propagule pressure and disturbance, which may facilitate exotic invasion. In contrast, disturbance from roads may increase the likelihood of local extinction for particular natives. Differing climatic preferences within the native and exotic species pools may also partially explain the observed patterns. There was no evidence that native diversity directly affects exotic diversity (or vice versa). Surprisingly, I found that connectivity was not an important predictor of richness or composition despite the high degree of habitat fragmentation in this ecosystem.

Many regional and local factors can influence the distribution of native and exotic species in ecological communities. I examined the regional- and local-scale determinants of native and exotic vascular plant species richness and composition in a highly fragmented oak savanna ecosystem on southeastern Vancouver Island. In sharp contrast to most reported results, I found a negative relationship between native and exotic richness at the regional scale, and no relationship at the local scale. Two extrinsic factors, surrounding road density and climate, best explained the regional-scale relationship by each affecting natives and exotics in opposite ways. Road density and climate were also the dominant predictors of native and exotic composition at the regional scale. Patterns in the patch occupancy of individual species confirmed the importance of these factors but I found that low surrounding road densities and cool, wet conditions predicted the presence of many natives and the absence of many exotics. Environmental factors explained variation in richness and composition at the local scale, but these factors were different for natives and exotics. My results suggest that natives and exotics respond to roads and climate in fundamentally different ways. Roads increase both exotic propagule pressure and disturbance, which may facilitate exotic invasion. In contrast, disturbance from roads may increase the likelihood of local extinction for particular natives. Differing climatic preferences within the native and exotic species pools may also partially explain the observed patterns. There was no evidence that native diversity directly affects exotic diversity (or vice versa). Surprisingly, I found that connectivity was not an important predictor of richness or composition despite the high degree of habitat fragmentation in this ecosystem. / Science, Faculty of / Botany, Department of / Graduate

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.