In order to predict future range shifts for invasive species it is important to explore their ability to acclimate to the new environment and understand physiological and reproductive constraints controlling their distribution. My dissertation studied mechanisms by which temperature may affect the distribution of two of the most aggressive plant invaders in North America, Bromus tectorum and Bromus rubens. While Bromus tectorum is dominant in the “cold desert” steppes of the Intermountain region of western North America, B. rubens is one of the severe grass invaders in the “hot deserts” of southwestern North America. I first evaluated whether winter freezing tolerance is the mechanism responsible for the distinct northern range limits of Bromus species. Bromus rubens has a slower rate of freezing acclimation that leads to intolerance of sudden, late-autumn reductions in temperature below -12°C, Bromus tectorum, by contrast, cold hardens rapidly and is not impacted by the sudden severe late-autumn cold. Photosynthetic response to temperature does not explain their current range separation. Bromus species differ little in their photosynthetic temperature responses and the acclimation pattern of photosynthesis. Both species acclimated to a broad range of temperature through the amelioration of Pi regeneration limitation at sub-optimal temperatures and improved carboxylation capacity above the thermal optimum which probably resulted from increased thermostability of Rubisco activase. The effect of elevated temperatures during flowering on the seed yield of Bromus species demonstrates that neither species produces seed at 36°C and above. These thresholds are close to temperatures encountered during flowering in their natural environment. In summary, climatic changes will cause northward range expansion of Bromus species due to less severe autumn and winter, while reproductive failure could cause range contraction at their southern margins.

With steady growth in global commerce and intensified ship traffic worldwide, comes the increased risk of invasion by non-indigenous organisms. Annually, >7000 vessels traveled across Galveston Bay, Texas from 2005-2010. These vessels discharged ~106 million metric tons of ballast water, equivalent to ~3.4% of the total volume of the Bay. A majority of these discharging vessels originated from around the Gulf of Mexico and the Caribbean Sea. By evaluating the source and frequency of inoculations from various locations, we are striving to assess the invasibility risk to Galveston Bay by way of ballast water. We identified organisms from Galveston Bay, ballast water samples and growout experiments using molecular methods. To our knowledge, this is the first utilization of molecular methods to identify the phytoplankton community within Galveston Bay. Within Galveston Bay, we identified 15 genera of dinoflagellates, 2 of which have previously gone undetected including Takayama and Woloszynskia. Thirteen ballast water samples yielded twenty genera of Protists, Fungi or Animalia from at least ten different phyla. With more than seven genera identified, dinoflagellates were the most diverse group: including the known toxin producer Pfiesteria and Scrippsiella which has not previously been detected in Galveston Bay. The most common diatoms in the ballast water samples were Actinocyclus, Ditylum, Nitzschia, Stephanopyxis and Thalassiosirales. At the termination of the growout experiments eight genera of phytoplankton were identified including: Dinophysis, Gymnodinium, Gyrodinium, Heterocapsa, Peridinium, Scrippsiella, Chaetoceros and Nitzschia. With these findings, Galveston Bay has the potential to be both a recipient and donor region of dinoflagellates. Dinoflagellates, capable of forming harmful algal blooms leading to fish and shellfish kills, are being transported to Galveston Bay via ballast water. Our results suggest that Galveston Bay is at risk for invasive species introductions via ballast water and support the idea that a monitoring system within the ports as well as the bay should be put in place. The actions would help to maintain the current health of this ecosystem and aide in preventing a negative impact in the event of successful establishment of a non-indigenous species of phytoplankton transported to Galveston Bay via ballast water.

Conservation in Canada is increasingly driven by land use planning processes. Approaches to governing nature conservation have shifted dramatically from protecting isolated pristine areas to greater attention to the remaining fragments of greenspace in urban, semi-urban and rural areas. The ways that societies govern and use nature are always changing, and these physical management actions are connected to deeply rooted cultural norms and values about the ideal relationship between humans and nature. In the land use planning approach to conservation, citizens and governments find value and construct meaning for remaining nature rather than beginning with normative considerations of what is most worthy of protection. At the root of this conservation planning trend is a growing appreciation for hybrid nature that is valued as natural in spite of the past or present influences upon it. This represents a dramatic shift from the traditional values involved in North American nature conservation, where nature was most valued for its perceived separation from human influence and protected to maintain its untouched qualities. In light of these ideological shifts in the ways that Canadians value and in turn manage nature, is there a corresponding change in the ways that conservation activists perceive environmental value and evaluate naturalness? An increasing number of studies demonstrate that public valuation of nature is not limited to pristine environments: even highly disturbed environments can be valued as natural and are not perceived as a form of lesser nature. Conceptions of what is natural and what is not are highly subjective and variable; in particular, the body of work on the social dimensions of both invasive species and ecological restoration demonstrates the ways in which people construct naturalness in accord with their values and cultural context. By exploring the extent to which people perceive invasive species as reducing naturalness and how ecological restoration is perceived to restore it, these subjects serve as excellent conceptual lenses for exploring constructions of nature. This study explores the subtle variations in environmental values and perception of naturalness among a study population who self-identify as pursuing the same goal: ensuring the continued protection of the Oak Ridges Moraine. The Moraine is a partly urbanized landform in southern Ontario that is situated within a complex hybrid socio-ecological landscape. It is also the subject of an active and high profile conservation movement that has spanned over 40 years. Using a combination of interviews and Q Method, this study explored how citizens engaged in Oak Ridges Moraine conservation perceive both the current and ideal state of naturalness on the Moraine, with specific emphasis on how the discourses these citizens use to frame the Moraine invoke the concept of naturalness Findings from this study reveal that Moraine activists represent a conservation paradox: they value the natural, non-human qualities of the landform, yet at the same time identify the Moraine as a hybrid landscape with both social and ecological qualities. In particular, respondents indicated a strong interest in naturalness in the context of invasive species and ecological restoration, yet at the same time identified the naturalness of the Moraine to be a lesser priority in the face of urban development pressures. In this way, citizens engaged in Moraine conservation respond to the hybrid quality of the Moraine landscape by moving beyond the binary distinction between nature and society, situating themselves as both apart from and a part of the landscape at the same time. This finding demonstrates how values for conserving nature are affected by hybridity between social and ecological systems, and suggests how embracing the paradox of hybrid nature can contribute to understanding and managing complex socio-ecological systems.

Russian thistle (Salsola tragus L.) invaded areas of native montane grassland important to winter survival of bighorn sheep (Ovis canadensis Shaw) were studied in Jasper National Park, Alberta, Canada. The biology of Russian thistle and its control in the Park were studied in the field and greenhouse. Russian thistle in grasslands were 9.1 cm tall with 37.5 seeds per plant, whereas larger plants in naturally disturbed habitats were 29.8 cm tall with 1562.4 seeds per plant. Plants travelled up to 4,180 m during dispersal. With soil seed contact, litter depth did not inhibit performance or survivability; without soil contact, thick litter reduced germination and plant performance. Russian thistle responded positively to increased greenhouse temperature and drier conditions. Seven control treatments involving herbicide, seeding mixes, hand pulling, and grazing exclusion were assessed. Grazing exclusion was the best field management option, increasing litter and biomass, while reducing Russian thistle density and biomass. / Land Reclamation and Remediation

Die vorliegende Arbeit beschäftigt sich mit der Analyse von Invasionsbiologie und ökologischem Verhalten sowie der Etablierung der gebietsfremden Baumart Fraxinus pennsylvanica in den Auenwäldern der mittleren Elbe (Sachsen-Anhalt). Fraxinus pennsylvanica wurde im 18. Jahrhundert aus NO-Amerika nach Europa und auch Deutschland eingeführt. Die Baumart hat heute im Untersuchungsgebiet in Waldflächen auf Auenstandorten einen Flächenanteil von 21,4 %. Aufgrund der hohen Stetigkeiten und einer fortschreitenden Ausbreitung von Fraxinus pennsylvanica ist eine Bedrohung der standortstypischen Zusammensetzung der Hartholzauenwälder nicht auszuschließen und stellt vor allem für Bereiche mit Prozessschutz ein Risiko dar. Anhand ausgewählter Entwicklungsstadien der Baumart beschreibt die vorliegende Arbeit das Ausbreitungs- und Etablierungsverhalten von Fraxinus pennsylvanica und hat das Ziel, die zu erwartende mögliche weitere Ausbreitung im naturschutzfachlichen Rahmen des Untersuchungsgebietes zu analysieren und zu bewerten. Die Arbeit gliedert sich in Untersuchungen zu anemochorer und hydrochorer Diasporenausbreitung, zum Keimungs-verhalten und zur Dormanz der Samen sowie zum Vorkommen der Verjüngung auf Standorten des Hartholzauenwaldes. Die Untersuchungen zur Invasionsbiologie und zum ökologischen Verhalten von Fraxinus pennsylvanica wollen einen Beitrag leisten, die bisherigen Wissenslücken zur gebietsfremden Baumart im neuen Areal zu schließen und Grundlagen sowie Anregungen für ein sinnvolles Managementkonzept liefern.

Caulerpa taxifolia is a marine alga notorious for its introduction and subsequent colonization of the Mediterranean Sea. It is recognized as one of the 100 worst invasive species, and it is suggested that much of its expansion may have been at the expense of native seagrass beds. To date, the bulk of research on C. taxifolia has centred on quantifying expansion and methods of eradication. There are few quantitative data on the relationship between C. taxifolia growth and environmental characteristics (e.g. light, temperature, nutrients). Furthermore, once C. taxifolia has been introduced to a system it is exceptionally difficult and expensive to eradicate. Accordingly the implications, both positive and negative, of this new habitat type must be considered in the context of the new habitat mosaic into which it fits. Australia is unique in that it has both native and invasive populations of C. taxifolia. These populations offer not only an opportunity to examine the dynamics of C. taxifolia beds and their associated communities at different latitudes and temperature and light regimes, but especially in the context of a high diversity marine coastal environment. The objectives of this thesis were to use native (Moreton Bay, Queensland) and invasive (Pittwater, New South Wales and Port River, South Australia) populations of C. taxifolia to: 1) quantify the relationship between environmental drivers (light, temperature, nutrients) and C. taxifolia growth, and 2) examine differences in habitat use and function between seagrass, C. taxifolia, unvegetated substrate. Most of the locations in Australia where large C. taxifolia beds occur are adjacent to urban areas that have a degraded water quality. Manipulative experiments in Moreton Bay demonstrated that nutrients stimulate C. taxifolia growth, however, light availability and seasonality appear to influence the response of C. taxifolia growth to nutrients. Short-term manipulative experiments were conducted across a range of seasons and locations, to capture the effects of temperature on growth. Temperature was the dominant factor affecting rate of stolon extension in both native and invasive locations. Colonization potential of C. taxifolia appears to be driven by ambient water quality (light and nutrients) and bed expansion is driven by temperature in systems where nutrients are saturating. Epifaunal communities sampled by beam trawl were dominated by the families Palaeomonidae, Terapontidae, Scorpaenidae, Monacanthidae, Syngnathidae, Gobiidae, and were largely similar between seagrasses and C. taxifolia; however, syngnathids were absent from C. taxifolia beds. I examined habitat use patterns between seagrass (Zostera muelleri), C. taxifolia, and unvegetated substrate. Fish preferred seagrass to C. taxifolia; however, in the absence of a seagrass fish used C. taxifolia. While C. taxifolia may have similar structural benefits to some seagrasses, there are fewer food resources available within C. taxifolia beds. Furthermore, grazing may be limited to a few specialist grazers. Within the habitat mosaic, C. taxifolia will provide some benefit over an unvegetated substrate; however, that benefit might mask losses in system quality or resilience by decreasing the threshold level for change within the community. Therefore, should a perturbation occur (e.g. sudden drop in water temperature, filamentous algal bloom) a system comprised solely of seagrass could withstand such stress; however, a habitat mosaic of seagrass and C. taxifolia could have a rapid and dramatic loss in its ability to sustain a diverse faunal community. Ultimately, it is most important to protect the system from anthropogenic degradation so it is more resilient to environmental changes.

Hieracium lepidulum is an invasive weed in New Zealand. It colonises a wide range of habitats including pine plantations, scrubland, native Nothofagus forest, and mid-altitude to alpine tussock grassland, where it is competing with indigenous species. Understanding the breeding systems and population genetic structure of H. lepidulum is important for biocontrol, and aids in the understanding of evolutionary colonisation processes. H. lepidulum is a triploid, diplosporous, obligate apomict. This type of reproduction through clonal seed does not involve meiosis or fertilisation, and theoretically populations should contain very low levels of genetic variation, the only source being somatic mutation. Common garden experiments and microsatellite markers were used to determine the population genetic structure of H. lepidulum populations in the Craigieburn Range, Canterbury. Both experiments revealed that populations, sampled from three replicate altitudes within three geographically-separated locations, contained no genetic variation; individuals all possessed the same microsatellite genotype. These results strongly suggest that the Craigieburn Range H. lepidulum individuals reproduce solely by apomixis and populations belong to the same clonal lineage. Populations were also examined for their response to two abiotic environmental ‘stresses’, drought and shade. H. lepidulum populations’ exhibited high drought tolerance, yet appeared to be shade-intolerant. Low levels of reproduction in light-limiting habitats will prevent the invasion of H. lepidulum into closed-canopy forest habitats. H. lepidulum appears to have overcome the reduction in fitness associated with apomictic reproduction by phenotypic plasticity, fixed heterozygosity and polyploidy – all associated with increased vigour, fitness, and the ability to occupy broader ecological niches. This study’s results are hopeful for the development of biocontrol programs involving genotype-specific pathogens but suggest that grazing management may not succeed. The data will be useful for future comparisons of genetic structure during the course of H. lepidulum invasions and will contribute to the management of this invasive weed.

Doctor of Philosphy (PhD) / This thesis describes the work carried out during four treks, each over 10-11 days, from 1400m to 5000m in the Nepal Himalaya and further work performed during several two-night sojourns at the Barcroft Laboratory at 3800m on White Mountain in California, USA. Nineteen volunteers were studied during the treks in Nepal and seven volunteers were studied at White Mountain. All subjects were normal, healthy individuals who had not travelled to altitudes higher than 1000m in the previous twelve months. The aims of this research were to examine the effects on sleep, and the ventilatory patterns during sleep, of incremental increases in altitude by employing portable polysomnography to measure and record physiological signals. A further aim of this research was to examine the relationship between the ventilatory responses to hypoxia and hypercapnia, measured at sea level, and the development of periodic breathing during sleep at high altitude. In the final part of this thesis the possibility of preventing and treating Acute Mountain Sickness with non-invasive positive pressure ventilation while sleeping at high altitude was tested. Chapter 1 describes the background information on sleep, and breathing during sleep, at high altitudes. Most of these studies were performed in hypobaric chambers to simulate various high altitudes. One study measured sleep at high altitude after trekking, but there are no studies which systematically measure sleep and breathing throughout the whole trek. Breathing during sleep at high altitude and the physiological elements of the control of breathing (under normal/sea level conditions and under the hypobaric, hypoxic conditions present at high altitude) are described in this Chapter. The occurrence of Acute Mountain Sickness (AMS) in subjects who travel form near sea level to altitudes above 3000m is common but its pathophysiology not well understood. The background research into AMS and its treatment and prevention are also covered in Chapter 1. Chapter 2 describes the equipment and methods used in this research, including the polysomnographic equipment used to record sleep and breathing at sea level and the high altitude locations, the portable blood gas analyser used in Nepal and the equipment and methodology used to measure each individual’s ventilatory response to hypoxia and hypercapnia at sea level before ascent to the high altitude locations. Chapter 3 reports the findings on the changes to sleep at high altitude, with particular focus on changes in the amounts of total sleep, the duration of each sleep stage and its percentage of total sleep, and the number and causes of arousals from sleep that occurred during sleep at increasing altitudes. The lightest stage of sleep, Stage 1 non-rapid eye movement (NREM) sleep, was increased, as expected with increases in altitude, while the deeper stages of sleep (Stages 3 and 4 NREM sleep, also called slow wave sleep), were decreased. The increase in Stage 1 NREM in this research is in agreement with all previous findings. However, slow wave sleep, although decreased, was present in most of our subjects at all altitudes in Nepal; this finding is in contrast to most previous work, which has found a very marked reduction, even absence, of slow wave sleep at high altitude. Surprisingly, unlike experimental animal studies of chronic hypoxia, REM sleep was well maintained at all altitudes. Stage 2 NREM and REM sleep, total sleep time, sleep efficiency and spontaneous arousals were maintained at near sea level values. The total arousal index was increased with increasing altitude and this was due to the increasing severity of periodic breathing as altitude increased. An interesting finding of this research was that fewer than half the periodic breathing apneas and hypopneas resulted in arousal from sleep. There was a minor degree of upper airway obstruction in some subjects at sea level but this was almost resolved by 3500m. Chapter 4 reports the findings on the effects on breathing during sleep of the progressive increase of altitude, in particular the occurrence of periodic breathing. This Chapter also reports the results of changes to arterial blood gases as subjects ascended to higher altitudes. As expected, arterial blood gases were markedly altered at even the lowest altitude in Nepal (1400m) and this change became more pronounced at each new, higher altitude. Most subjects developed periodic breathing at high altitude but there was a wide variability between subjects as well as variability in the degree of periodic breathing that individual subjects developed at different altitudes. Some subjects developed periodic breathing at even the lowest altitude and this increased with increasing altitude; other subjects developed periodic breathing at one or two altitudes, while four subjects did not develop periodic breathing at any altitude. Ventilatory responses to hypoxia and hypercapnia, measured at sea level before departure to high altitude, was not significantly related to the development of periodic breathing when the group was analysed as a whole. However, when the subjects were grouped according to the steepness of their ventilatory response slopes, there was a pattern of higher amounts of periodic breathing in subjects with steeper ventilatory responses. Chapter 5 reports the findings of an experimental study carried out in the University of California, San Diego, Barcroft Laboratory on White Mountain in California. Seven subjects drove from sea level to 3800m in one day and stayed at this altitude for two nights. On one of the nights the subjects slept using a non-invasive positive pressure device via a face mask and this was found to significantly improve the sleeping oxyhemoglobin saturation. The use of the device was also found to eliminate the symptoms of Acute Mountain Sickness, as measured by the Lake Louise scoring system. This finding appears to confirm the hypothesis that lower oxygen saturation, particularly during sleep, is strongly correlated to the development of Acute Mountain Sickness and may represent a new treatment and prevention strategy for this very common high altitude disorder.

Diss. (sammanfattning) Stockholm : Karol. inst., 2003. / Härtill 4 uppsatser.

Thesis (M.Sc.) - Carleton University, 2006. / Includes bibliographical references (p. 37-43). Also available in electronic format on the Internet.