Many recent studies have demonstrated that propagule pressure is a useful predictor of patterns of invasions by non-native species. However, most of these studies have used only current, not historical, data to estimate propagule pressure. Recognizing the potential importance of propagule pressure over time, I used surrogate variables that represent both past and present propagule pressure, for example, the length of time a surrounding area had been developed. I quantified the relationships between these surrogate variables and the distribution and abundance of non-native woody plant species in central Texas. I constructed statistical models predicting native and non-native species richness and the occurrence of five common species using a set of six ecological and five development-related predictor variables. I compared all models using the corrected Akaike information criterion (AICc). Overall, age of residential development surrounding native woodlands was the best predictor, other than community type, of non-native species richness. As expected, areas near older developments had more non-native species than areas near newer developments. Surprisingly, age of development and average city age, two different measures of the length of time that landscaping (a major source of propagules of non-native woody species in this region) had been present nearby, were much better predictors than distance to source populations. Age of development and average city age (weighted by distance from the site) were also both correlated with distance to source populations; this may be true in other systems as well. This suggests that the reason distance to source population has been a successful predictor of invasion may be because it is a surrogate for an underlying causal variable, length of time of exposure to source populations. Future studies of non-native invasions would benefit from taking into account both past and present propagule pressure: age of residential development and city age could be useful surrogates in other systems. / text

No description available.

The spread of invasive species poses a major ecological and economical threat. Consequently there are ongoing efforts to develop a generalizable mechanism to predict establishment success of non-native species. One proposed mechanism to predict establishment success is propagule pressure, which is defined as the number of individuals introduced at a given time. Although some studies have demonstrated a positive correlation between propagule pressure and establishment success, others have not, and the effect of propagule pressure on establishment success remains unclear. To address this challenge, a strain of bacteria engineered with an Allee effect, a growth dynamic that is often associated with establishing species, was used. The timing between successive introduction events that resulted in establishment success was measured. It was observed that if the time between two introduction events was sufficiently long, growth did not occur. By manipulating the growth rate of the bacteria, it was shown that that the minimal time between the two introduction events that resulted in growth was constrained as growth rate decreased. Moreover, it was concluded that asymmetry in the density of bacteria introduced in the introduction events increased the maximum time between introduction events that resulted in growth. These results help to remedy conflicting data in the literature by identifying conditions where propagule pressure has, and does not have, a positive impact on establishment success. These findings can have major implications in understanding and predicting the unique population dynamics of invasive species.

Doctor of Philosophy (PhD) / Early recognition of plant invaders is key to their successful management. Yet knowledge of the ecological impacts of species before they become widespread is poor. This thesis examines the ecology of invasive Pinus radiata, a species which is known to spread from introduced plantings in Australia but is currently a low profile invader. Pinus invasions are considered major ecological problems in New Zealand and South Africa where wildlings are beginning to dominate natural areas and suppress native vegetation. Invasion success elsewhere and the large softwood estate in Australia suggest that pines may begin to dominate native eucalypt forests bordering large commercial plantations. This research focused on three components of impact of P. radiata; extent, abundance and effect per individual. The borders of 29 P. radiata plantations in NSW were surveyed in order to quantify the current level of invasion and to identify factors facilitating pine spread. Of particular interest was the role of propagule pressure, vegetation type and fire in the invasion process. The area of land in NSW currently invaded by P. radiata was estimated at almost 4 500 ha, although this is likely to be an underestimate due to an inability to detect wildlings (self-sown pines) at long distances from the plantation. Twenty six of the 29 plantations produced wild pines, however most of the sites are in the very early stages of invasion. Noticeable wildling populations were recorded at nine sites indicating that P. radiata is capable of establishing within native vegetation. Pine spread was most severe in the world heritage listed Blue Mountains region where pine densities reached up to 2000 per hectare in areas adjacent to the plantation and isolated pines were recorded up to 4 km from the source. The presence of isolated pines within intact native vegetation suggests that disturbance is not required for pine establishment in forested environments. Furthermore, high pine emergence and survival rates in eucalypt woodland and evidence of self reproduction by wildlings suggest that in the absence of adequate control measures pines may become established invaders in the Australian landscape. While low levels of current invasion at many sites hindered the ability to examine the factors facilitating invasion some variables that appear to be driving pine success were identified. At the landscape scale plantation size and residence time were significant predictors of the level of invasion at a site. Areas of native vegetation vi adjacent to plantations less than 40 years experienced very low levels of invasion suggesting a lag period between plantation establishment and invasion. However, pines with diameters up to 60 cm were observed growing adjacent to plantations younger than 40 years implying that the first colonisers are capable of establishing soon after plantation trees become reproductive. Propagule pressure was also found to have a strong influence on invasion success on a smaller scale manifesting in a significant positive relationship between the age of a plantation compartment and the likelihood of invasion. A negative relationship between plantation size and level of invasion was a surprising result and was influenced by just two large sites that happened to be located in areas of high rainfall. All sites receiving more than 1300 mm annual rainfall experienced low levels of invasion suggesting that this is a limiting factor for pine spread in NSW. There were significant differences in the level of invasion between vegetation types implying that some communities are more susceptible to invasion. Patterns of spread confirmed ideas regarding the facilitative effect of disturbance in the invasion process and the resistance of wet sclerophyll forest to invasion in Australia. An absence of wildlings in cleared land and areas of remnant bushland was attributed to high levels of grazing pressure. Wind direction did not appear to influence the distribution of pines close to the plantation, but evidence of long distance wind dispersal of pines was provided by an investigation of pine spread from the air at one site where large pines were found growing 10 km downwind from a mature plantation. Fire was found to have both a positive and negative influence on the invasion process. High intensity wildfires are capable of destroying large pines with diameters exceeding 50 cm. However, fire can stimulate seed release from cones resulting in large post-fire recruitment pulses. Seedling densities of up to 3050 per hectare were recorded almost 3 years after wildfire, suggesting that follow up control prior to recruits reaching coning age, i.e. within 5 years, would be beneficial. Surveys of wildling pines exposed to low intensity hazard reduction burns suggest that the majority of pines greater than 3 m in height and with a diameter of more than 10 cm will survive the fire. Low intensity prescribed fires that are carried out after pines have reached this size will fail to control wildling populations. To examine the influence of P. radiata once it has established in the native community this study focused on two mechanisms of impact, the addition of pine litter and increased shade due to an increase in canopy cover. Collection of pine litterfall vii within an invaded eucalypt woodland over a 2 year period recorded rates of up to 1400 kg/ha/year in the most heavily invaded area with a pine basal area of 11.3m2/ha. More than 70 % of pine litter fell directly below the pine canopy suggesting that the most severe litter effects will be limited to these areas. Glasshouse and field experiments were conducted to examine the influence of this increased litter load on the emergence of P. radiata and two native species. Recruitment of native plant species was impeded by litter levels of 6000 kg/ha, the equivalent of approximately 4 years of pine litterfall. Both P. radiata and the two natives responded similarly to pine and eucalypt litter suggesting the two litter types are influencing the recruitment phase equally. However, where pines are added to the system, increased litterfall rates could potentially result in the doubling of the litter load and hence a greater barrier to seedling establishment. Pine invaded eucalypt woodlands are also subjected to three fold increases in canopy cover. Trends in reduced emergence of native species under a pine canopy suggest that the addition of pines to eucalypt forests is likely to have a negative influence on native recruitment and may result in a shift towards a shade tolerant community. However, reversal of trends in emergence below pine canopy between seasons implies that quantifying invasion impacts requires a consideration of temporal variation. Increased levels of disturbance, forest fragmentation and an increasing pine estate are likely to lead to the infestation of new areas. Furthermore the lag phase associated with pine spread means that even if no new plantations are established the number of invasion events will increase. This study has identified a number of risk factors that can be used to guide plantation establishment and the management of invasion events. Minimising disturbance at plantation borders and increasing the ‘no planting’ zone will help to reduce the impacts of pines. Where possible new plantations should be established upwind of cleared land or at least, wet sclerophyll forest. Frequent monitoring of the borders of plantations yet to source invasions, particularly those greater than 40 years of age, will help identify problem areas before control becomes difficult and costly. Maps of the 29 plantations marked with areas of pine infestation will help prioritise sites for control and provide base level knowledge for future monitoring of pine spread. Stringent legislation that binds plantation managers to control wildlings beyond their boundaries is critical for the effective management of pine invasions. With infinite numbers of invaders and limited funds to dedicate to their control, a method of triaging species for management is critical. This is particularly difficult viii when information is typically biased towards invaders that are already widespread. By focussing on the ecological impacts of invaders it becomes possible to rank species on the basis of the threat they pose to native communities. Ecological research is capable of providing the knowledge to quantify invasion impacts and must remain at the centre of policy decisions.

The global movement of aquatic non-indigenous species can have severe ecological, environmental and economic impacts emphasizing the need to identify potential invaders and transport pathways. Initial transport is arguably the most important stage of the invasion process owing to its role in selectively determining potential invasion candidates. This study characterizes a well defined human-mediated dispersal mechanism, ballast water transport, as a vector for the introduction of non-indigenous zooplankton. Ballast water exchange in the open ocean is the most widely adopted practice for reducing the threat of aquatic invasions and is mandatory for most foreign vessels intending to release ballast in Canadian waters. Ships entering Canadian ports are categorized into the following three shipping classes based on current regulations: overseas vessels carrying exchanged ballast water, intra-coastal vessels carrying exchanged ballast water or intra-coastal vessels carrying un-exchanged ballast water. This study characterizes zooplankton communities associated with each of these shipping classes sampled from ports on Canada’s Pacific coast, Atlantic coast and the Great Lakes Basin. Ballast water samples were collected and analyzed from 77 vessels between 2006 - 2007. The ballast water environment was found to be diverse, with over 193 zooplankton taxa, 71 of which were non-indigenous to their receiving environments. Intracoastal vessels containing un-exchanged coastal water transported the greatest density of non-indigenous zooplankton into Canadian ports. Total zooplankton density was found to be negatively correlated with ballast water age The absence of mandatory ballast water exchange and the younger ballast water age of coastal un-exchanged vessels is likely responsible for the higher density of non-indigenous zooplankton in intracoastal un-exchanged vessels. Propagule pressure, invasion history and environmental suitability are all useful in evaluating invasion potential and all suggest that intracoastal un-exchanged vessels pose the greatest invasion threat to Canadian aquatic ecosystems. In conclusion, although the risk of primary introductions from overseas ports may have been reduced through open-ocean exchange of ballast water, secondary introductions from previously invaded ports in North America may be the primary threat to Canadian aquatic ecosystems via this transport vector.

The global movement of aquatic non-indigenous species can have severe ecological, environmental and economic impacts emphasizing the need to identify potential invaders and transport pathways. Initial transport is arguably the most important stage of the invasion process owing to its role in selectively determining potential invasion candidates. This study characterizes a well defined human-mediated dispersal mechanism, ballast water transport, as a vector for the introduction of non-indigenous zooplankton. Ballast water exchange in the open ocean is the most widely adopted practice for reducing the threat of aquatic invasions and is mandatory for most foreign vessels intending to release ballast in Canadian waters. Ships entering Canadian ports are categorized into the following three shipping classes based on current regulations: overseas vessels carrying exchanged ballast water, intra-coastal vessels carrying exchanged ballast water or intra-coastal vessels carrying un-exchanged ballast water. This study characterizes zooplankton communities associated with each of these shipping classes sampled from ports on Canada’s Pacific coast, Atlantic coast and the Great Lakes Basin. Ballast water samples were collected and analyzed from 77 vessels between 2006 - 2007. The ballast water environment was found to be diverse, with over 193 zooplankton taxa, 71 of which were non-indigenous to their receiving environments. Intracoastal vessels containing un-exchanged coastal water transported the greatest density of non-indigenous zooplankton into Canadian ports. Total zooplankton density was found to be negatively correlated with ballast water age The absence of mandatory ballast water exchange and the younger ballast water age of coastal un-exchanged vessels is likely responsible for the higher density of non-indigenous zooplankton in intracoastal un-exchanged vessels. Propagule pressure, invasion history and environmental suitability are all useful in evaluating invasion potential and all suggest that intracoastal un-exchanged vessels pose the greatest invasion threat to Canadian aquatic ecosystems. In conclusion, although the risk of primary introductions from overseas ports may have been reduced through open-ocean exchange of ballast water, secondary introductions from previously invaded ports in North America may be the primary threat to Canadian aquatic ecosystems via this transport vector.

Doctor of Philosophy (PhD) / Early recognition of plant invaders is key to their successful management. Yet knowledge of the ecological impacts of species before they become widespread is poor. This thesis examines the ecology of invasive Pinus radiata, a species which is known to spread from introduced plantings in Australia but is currently a low profile invader. Pinus invasions are considered major ecological problems in New Zealand and South Africa where wildlings are beginning to dominate natural areas and suppress native vegetation. Invasion success elsewhere and the large softwood estate in Australia suggest that pines may begin to dominate native eucalypt forests bordering large commercial plantations. This research focused on three components of impact of P. radiata; extent, abundance and effect per individual. The borders of 29 P. radiata plantations in NSW were surveyed in order to quantify the current level of invasion and to identify factors facilitating pine spread. Of particular interest was the role of propagule pressure, vegetation type and fire in the invasion process. The area of land in NSW currently invaded by P. radiata was estimated at almost 4 500 ha, although this is likely to be an underestimate due to an inability to detect wildlings (self-sown pines) at long distances from the plantation. Twenty six of the 29 plantations produced wild pines, however most of the sites are in the very early stages of invasion. Noticeable wildling populations were recorded at nine sites indicating that P. radiata is capable of establishing within native vegetation. Pine spread was most severe in the world heritage listed Blue Mountains region where pine densities reached up to 2000 per hectare in areas adjacent to the plantation and isolated pines were recorded up to 4 km from the source. The presence of isolated pines within intact native vegetation suggests that disturbance is not required for pine establishment in forested environments. Furthermore, high pine emergence and survival rates in eucalypt woodland and evidence of self reproduction by wildlings suggest that in the absence of adequate control measures pines may become established invaders in the Australian landscape. While low levels of current invasion at many sites hindered the ability to examine the factors facilitating invasion some variables that appear to be driving pine success were identified. At the landscape scale plantation size and residence time were significant predictors of the level of invasion at a site. Areas of native vegetation vi adjacent to plantations less than 40 years experienced very low levels of invasion suggesting a lag period between plantation establishment and invasion. However, pines with diameters up to 60 cm were observed growing adjacent to plantations younger than 40 years implying that the first colonisers are capable of establishing soon after plantation trees become reproductive. Propagule pressure was also found to have a strong influence on invasion success on a smaller scale manifesting in a significant positive relationship between the age of a plantation compartment and the likelihood of invasion. A negative relationship between plantation size and level of invasion was a surprising result and was influenced by just two large sites that happened to be located in areas of high rainfall. All sites receiving more than 1300 mm annual rainfall experienced low levels of invasion suggesting that this is a limiting factor for pine spread in NSW. There were significant differences in the level of invasion between vegetation types implying that some communities are more susceptible to invasion. Patterns of spread confirmed ideas regarding the facilitative effect of disturbance in the invasion process and the resistance of wet sclerophyll forest to invasion in Australia. An absence of wildlings in cleared land and areas of remnant bushland was attributed to high levels of grazing pressure. Wind direction did not appear to influence the distribution of pines close to the plantation, but evidence of long distance wind dispersal of pines was provided by an investigation of pine spread from the air at one site where large pines were found growing 10 km downwind from a mature plantation. Fire was found to have both a positive and negative influence on the invasion process. High intensity wildfires are capable of destroying large pines with diameters exceeding 50 cm. However, fire can stimulate seed release from cones resulting in large post-fire recruitment pulses. Seedling densities of up to 3050 per hectare were recorded almost 3 years after wildfire, suggesting that follow up control prior to recruits reaching coning age, i.e. within 5 years, would be beneficial. Surveys of wildling pines exposed to low intensity hazard reduction burns suggest that the majority of pines greater than 3 m in height and with a diameter of more than 10 cm will survive the fire. Low intensity prescribed fires that are carried out after pines have reached this size will fail to control wildling populations. To examine the influence of P. radiata once it has established in the native community this study focused on two mechanisms of impact, the addition of pine litter and increased shade due to an increase in canopy cover. Collection of pine litterfall vii within an invaded eucalypt woodland over a 2 year period recorded rates of up to 1400 kg/ha/year in the most heavily invaded area with a pine basal area of 11.3m2/ha. More than 70 % of pine litter fell directly below the pine canopy suggesting that the most severe litter effects will be limited to these areas. Glasshouse and field experiments were conducted to examine the influence of this increased litter load on the emergence of P. radiata and two native species. Recruitment of native plant species was impeded by litter levels of 6000 kg/ha, the equivalent of approximately 4 years of pine litterfall. Both P. radiata and the two natives responded similarly to pine and eucalypt litter suggesting the two litter types are influencing the recruitment phase equally. However, where pines are added to the system, increased litterfall rates could potentially result in the doubling of the litter load and hence a greater barrier to seedling establishment. Pine invaded eucalypt woodlands are also subjected to three fold increases in canopy cover. Trends in reduced emergence of native species under a pine canopy suggest that the addition of pines to eucalypt forests is likely to have a negative influence on native recruitment and may result in a shift towards a shade tolerant community. However, reversal of trends in emergence below pine canopy between seasons implies that quantifying invasion impacts requires a consideration of temporal variation. Increased levels of disturbance, forest fragmentation and an increasing pine estate are likely to lead to the infestation of new areas. Furthermore the lag phase associated with pine spread means that even if no new plantations are established the number of invasion events will increase. This study has identified a number of risk factors that can be used to guide plantation establishment and the management of invasion events. Minimising disturbance at plantation borders and increasing the ‘no planting’ zone will help to reduce the impacts of pines. Where possible new plantations should be established upwind of cleared land or at least, wet sclerophyll forest. Frequent monitoring of the borders of plantations yet to source invasions, particularly those greater than 40 years of age, will help identify problem areas before control becomes difficult and costly. Maps of the 29 plantations marked with areas of pine infestation will help prioritise sites for control and provide base level knowledge for future monitoring of pine spread. Stringent legislation that binds plantation managers to control wildlings beyond their boundaries is critical for the effective management of pine invasions. With infinite numbers of invaders and limited funds to dedicate to their control, a method of triaging species for management is critical. This is particularly difficult viii when information is typically biased towards invaders that are already widespread. By focussing on the ecological impacts of invaders it becomes possible to rank species on the basis of the threat they pose to native communities. Ecological research is capable of providing the knowledge to quantify invasion impacts and must remain at the centre of policy decisions.

The global movement of aquatic non-indigenous species can have severe ecological, environmental and economic impacts emphasizing the need to identify potential invaders and transport pathways. Initial transport is arguably the most important stage of the invasion process owing to its role in selectively determining potential invasion candidates. This study characterizes a well defined human-mediated dispersal mechanism, ballast water transport, as a vector for the introduction of non-indigenous zooplankton. Ballast water exchange in the open ocean is the most widely adopted practice for reducing the threat of aquatic invasions and is mandatory for most foreign vessels intending to release ballast in Canadian waters. Ships entering Canadian ports are categorized into the following three shipping classes based on current regulations: overseas vessels carrying exchanged ballast water, intra-coastal vessels carrying exchanged ballast water or intra-coastal vessels carrying un-exchanged ballast water. This study characterizes zooplankton communities associated with each of these shipping classes sampled from ports on Canada’s Pacific coast, Atlantic coast and the Great Lakes Basin. Ballast water samples were collected and analyzed from 77 vessels between 2006 - 2007. The ballast water environment was found to be diverse, with over 193 zooplankton taxa, 71 of which were non-indigenous to their receiving environments. Intracoastal vessels containing un-exchanged coastal water transported the greatest density of non-indigenous zooplankton into Canadian ports. Total zooplankton density was found to be negatively correlated with ballast water age The absence of mandatory ballast water exchange and the younger ballast water age of coastal un-exchanged vessels is likely responsible for the higher density of non-indigenous zooplankton in intracoastal un-exchanged vessels. Propagule pressure, invasion history and environmental suitability are all useful in evaluating invasion potential and all suggest that intracoastal un-exchanged vessels pose the greatest invasion threat to Canadian aquatic ecosystems. In conclusion, although the risk of primary introductions from overseas ports may have been reduced through open-ocean exchange of ballast water, secondary introductions from previously invaded ports in North America may be the primary threat to Canadian aquatic ecosystems via this transport vector. / Science, Faculty of / Earth, Ocean and Atmospheric Sciences, Department of / Graduate

Over the last century, flowering rush (Butomus umbellatus L.: Butomaceae) has escaped its native Eurasian range and has become a problematic species in North America. As an aquatic invasive species, flowering rush has degraded native wetlands and has interfered with human water usage. Although experimental work has been done regarding the reproductive biology of the species, few empirical studies regarding the ecology of the species have been conducted. The research reported here demonstrates that flowering rush is capable of aggressive clonal growth and propagation, and can perform well along a depth gradient from zero to 132cm. Proper management and control of invasive species relies on sound ecological knowledge of the target species, and this work aims to help gather that information.

Biological invasions are a significant area of research due to perceived negative environmental and economic impacts. The study of biological invasions has identified three broad components involved in successful invasions: propagule pressure, abiotic and biotic conditions. Propagule pressure is the product of introduced propagules and the frequency of those introductions, and is considered a driver in all stages of invasion. Data to quantify propagule pressure is often unavailable, and therefore, this research approached genetic information to produce estimates of propagule pressure in the successful invasion of cogongrass (Imperata cylindrica) in the United States. The following research utilized molecular methodologies to estimate genetic diversity and to infer historical introductions. Population genetic analyses were conducted for the purpose of estimating extant population-level genetic diversity at multiple-scales to first address documented cogongrass introduction(s) into this country, followed by an exploration of substantial range expansion into seven states across the region. Lastly, this research explicitly seeks evidence to support interspecific hybridization between cogongrass and a co-occurring congeneric (Imperata brasiliensis) having occurred. Aggressive range expansion by cogongrass has been attributed to such a hybridization event, but has not yet been specifically tested. Historical accounts of foreign introduction of cogongrass propagules identify two separate introductions of distinct source material made into Mississippi and Alabama. Localized population genetic analysis of these two states found substantial genetic variability within and among cogongrass populations, and supported a two-introduction scenario of distinct genetic source propagules establishing and subsequently intermixing. Enlarging the geographic scale of study incorporated five additional U.S. states currently experiencing and managing cogongrass invasion. Considerable genetic variability was found within and among the seven states surveyed. It was found that range expansion was unequal across the range, and that the most distant states tested were not genetically isolated from source populations, suggesting a possible anthropogenic role. Focusing solely on Florida where two congeners overlap ranges, morphology, and ecology, this population genetic analysis failed to detect significant evidence to support interspecific hybridization. Collectively, these investigations explored genome-level dynamics during invasion by a noteworthy invasive grass in an effort to better understand the process of biological invasions.