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• 281. [Article] Understanding the Importance of Intermittently Fragmented Stream Habitat for Isolated Westslope Cutthroat Trout (<i>Oncorhynchus clarki lewisi</i>) in the Colville National Forest, Washington

  Climate change and anthropogenic effects have vastly reduced Westslope Cutthroat Trout (Oncorhynchus clarki lewisi, WCT) habitat throughout their range, including the Colville National Forest in northeastern ...

  Citation × Citation Citation Abstract Copy Title: Understanding the Importance of Intermittently Fragmented Stream Habitat for Isolated Westslope Cutthroat Trout (<i>Oncorhynchus clarki lewisi</i>) in the Colville National Forest, Washington Author: Carpenter, Forrest Michael Year: 2016 Climate change and anthropogenic effects have vastly reduced Westslope Cutthroat Trout (Oncorhynchus clarki lewisi, WCT) habitat throughout their range, including the Colville National Forest in northeastern Washington where this study was conducted. Many native salmonid populations have declined in abundance since the early 1900s due to a variety of climate- and human-driven forces. Westslope Cutthroat Trout are especially sensitive to habitat loss or degradation and to climate change. Together, climate change, habitat degradation, and non-native salmonid invasions are contributing to increasingly fragmented WCT populations. Ongoing and predicted future warming trends are expected to further fragment these populations and isolate them in headwater stream reaches, with populations in the spatial margins of their distributions facing greater risk. Native salmonid populations are often separated or isolated by natural or artificial upstream migration barriers (i.e., waterfalls, culverts, etc.). Prior to continuing conservation and management actions targeting WCT, it is imperative to understand habitat requirements of this keystone species in fragmented areas. Field survey data were collected in the summer of 2015 on channel geomorphic characteristics and WCT presence/absence in 26 streams located in the Colville National Forest. A clear spatial separation was observed between Eastern Brook Trout (Salvenius fontinalis, EBT) and WCT above four culvert road crossings and the habitat in both of these areas was compared statistically to identify explicit differences. This dataset was also analyzed using logistic regression modeling to determine the best habitat predictors of the presence of isolated WCT populations existing upstream of these crossing. In general, stream habitat in the Middle and South Forks of Mill Creek had low large substrate, high fine sediments, and exhibited pool-riffle channel morphology. Pool habitat supporting isolated WCT was significantly smaller, in terms of volume and surface area, than pool habitat supporting sympatric populations of WCT and EBT, largely due to the headwater nature of channel units supporting isolated WCT populations. Additionally, due to the extreme drought conditions during 2015, stream flow was substantially diminished in the study area causing these reaches to be highly fragmented and largely disconnected from the rest of the stream channel. Fine sediments were generally higher in headwater reaches supporting isolated WCT, including in pools and riffles, which was unexpected, mainly because they exist above sediment delivery points in the longitudinal extent of the system. Logistic regression analysis indicated that the presence of isolated WCT populations was primarily positively associated with an increase in large wood and boulders, and negatively associated with increasing gravel, bedrock, habitat unit length, depth, and width (Significant x2, R2=0.174, misclassification rate = 14.9%, α=0.05). The final model correctly predicted 37.5% of isolated WCT presence observations and 96.5% of the WCT/EBT presence observations significantly better than by chance alone (k=0.81). This model, in fact, may be useful in identifying limited habitat due to the fragmented nature of the channel units supporting IWCT. Large wood and boulders were positively correlated to WCT presence, likely because both are important in the formation of pools and cascades. Channel unit length, width, depth, active channel width as well as gravel and bedrock substrates, were all negatively associated with WCT presence. This suggests that isolated WCT are primarily associated with small headwater cascades with complex shelter, which may provide greater thermal and predation refuge compared to shallow glide or large pool habitats. Future model analysis should include additional habitat variables such as water temperature, stream gradient, and species interactions to strengthen the prediction of Westslope Cutthroat Trout presence. Overall, I concluded that differences in stream habitat above and below blocking culverts are not driving Westslope Cutthroat Trout distributions in the study area due to confounding factors such as the presence of problematic barriers and small sample size. I also conclude that future conservation and management decisions specific to WCT should prioritize complex cascade habitat in headwater stream reaches because of the type and quantity of habitat they may provide, especially during severe drought or low flow conditions. Title: Understanding the Importance of Intermittently Fragmented Stream Habitat for Isolated Westslope Cutthroat Trout (<i>Oncorhynchus clarki lewisi</i>) in the Colville National Forest, Washington Author: Carpenter, Forrest Michael Year: 2016 Climate change and anthropogenic effects have vastly reduced Westslope Cutthroat Trout (Oncorhynchus clarki lewisi, WCT) habitat throughout their range, including the Colville National Forest in northeastern Washington where this study was conducted. Many native salmonid populations have declined in abundance since the early 1900s due to a variety of climate- and human-driven forces. Westslope Cutthroat Trout are especially sensitive to habitat loss or degradation and to climate change. Together, climate change, habitat degradation, and non-native salmonid invasions are contributing to increasingly fragmented WCT populations. Ongoing and predicted future warming trends are expected to further fragment these populations and isolate them in headwater stream reaches, with populations in the spatial margins of their distributions facing greater risk. Native salmonid populations are often separated or isolated by natural or artificial upstream migration barriers (i.e., waterfalls, culverts, etc.). Prior to continuing conservation and management actions targeting WCT, it is imperative to understand habitat requirements of this keystone species in fragmented areas. Field survey data were collected in the summer of 2015 on channel geomorphic characteristics and WCT presence/absence in 26 streams located in the Colville National Forest. A clear spatial separation was observed between Eastern Brook Trout (Salvenius fontinalis, EBT) and WCT above four culvert road crossings and the habitat in both of these areas was compared statistically to identify explicit differences. This dataset was also analyzed using logistic regression modeling to determine the best habitat predictors of the presence of isolated WCT populations existing upstream of these crossing. In general, stream habitat in the Middle and South Forks of Mill Creek had low large substrate, high fine sediments, and exhibited pool-riffle channel morphology. Pool habitat supporting isolated WCT was significantly smaller, in terms of volume and surface area, than pool habitat supporting sympatric populations of WCT and EBT, largely due to the headwater nature of channel units supporting isolated WCT populations. Additionally, due to the extreme drought conditions during 2015, stream flow was substantially diminished in the study area causing these reaches to be highly fragmented and largely disconnected from the rest of the stream channel. Fine sediments were generally higher in headwater reaches supporting isolated WCT, including in pools and riffles, which was unexpected, mainly because they exist above sediment delivery points in the longitudinal extent of the system. Logistic regression analysis indicated that the presence of isolated WCT populations was primarily positively associated with an increase in large wood and boulders, and negatively associated with increasing gravel, bedrock, habitat unit length, depth, and width (Significant x2, R2=0.174, misclassification rate = 14.9%, α=0.05). The final model correctly predicted 37.5% of isolated WCT presence observations and 96.5% of the WCT/EBT presence observations significantly better than by chance alone (k=0.81). This model, in fact, may be useful in identifying limited habitat due to the fragmented nature of the channel units supporting IWCT. Large wood and boulders were positively correlated to WCT presence, likely because both are important in the formation of pools and cascades. Channel unit length, width, depth, active channel width as well as gravel and bedrock substrates, were all negatively associated with WCT presence. This suggests that isolated WCT are primarily associated with small headwater cascades with complex shelter, which may provide greater thermal and predation refuge compared to shallow glide or large pool habitats. Future model analysis should include additional habitat variables such as water temperature, stream gradient, and species interactions to strengthen the prediction of Westslope Cutthroat Trout presence. Overall, I concluded that differences in stream habitat above and below blocking culverts are not driving Westslope Cutthroat Trout distributions in the study area due to confounding factors such as the presence of problematic barriers and small sample size. I also conclude that future conservation and management decisions specific to WCT should prioritize complex cascade habitat in headwater stream reaches because of the type and quantity of habitat they may provide, especially during severe drought or low flow conditions. Close

• 282. [Article] Examination of Human Impacts on the Biodiversity and Ecology of Lichen and Moss Communities

  Globally, more than half of the world's population is living in urban areas and it is well accepted that human activities (e.g. climate warming, pollution, landscape homogenization) pose a multitude of ...

  Citation × Citation Citation Abstract Copy Title: Examination of Human Impacts on the Biodiversity and Ecology of Lichen and Moss Communities Author: Prather, Hannah Marie Year: 2017 Globally, more than half of the world's population is living in urban areas and it is well accepted that human activities (e.g. climate warming, pollution, landscape homogenization) pose a multitude of threats to ecosystems. Largely, human-related impacts on biodiversity will hold consequences for larger ecological processes and research looking into human impacts on sensitive epiphytic lichen and moss communities is an emerging area of research. While seemingly small, lichen and moss communities exist on nearly every terrestrial ecosystem on Earth and contribute to whole-system processes (e.g. hydrology, mineral cycling, food web energetics) worldwide. To further examine human impacts on epiphytic communities, I conducted three studies examining urbanization and climate warming effects on epiphytic lichen and moss biodiversity and ecology. In the first study I revisited a historic urban lichen community study to assess how urban lichen communities have responded to regional air quality changes occurring over the last nearly two decades. I further investigated, for the first time, the biodiversity of urban tree canopy-dwelling lichen communities in a native coniferous tree species, Pseudotsuga menziesii. I found that urban parks and forested areas harbor a species rich community of lichens epiphytes. Further, I found evidence for the distinct homogenization of urban epiphytic lichen communities, suggesting that expanding beyond simplistic measures of biodiversity to consider community composition and functional biodiversity may be necessary when assessing the ecology and potential ecosystem services of epiphyte communites within urbanizing landscapes. Next, I present the first tall tree canopy study across a regional gradient of urbanization near Portland, Oregon, USA. I found that tall tree canopy epiphyte communities change dramatically along gradients of increasing urbanization, most notably by the transitioning of species functional groups from sensitive, oligotrophic species to a dominance of urban-tolerant, eutrophic species. The implications these dramatic shifts in species composition have on essential PNW ecosystem processes, like N-fixation and canopy microclimate regulation, is still not well understood and is difficult to formally evaluate. However, I find strong evidence that native conifer trees in urban areas may provide a diversity of essential ecosystem services, including providing stratified habitat for epiphyte communities and their associated micro arthropod communities and the scavenging of atmospherically deposited nutrients. Future work is needed to understand how losses in canopy N fixation and species with large biomass (both lichens and bryophytes) will affect nutrient and hydrologic cycling in the PNW region, which continue to undergo rapid growth and urbanization. The final chapter investigates the impacts of passive warming by Open Top Chambers (OTCs) in moss-dominated ecosystems located on the Western Antarctic Peninsula, an area of increasing climate warming. I compared species-specific temperature effects, moss canopy morphology, sexual reproductive effort and invertebrate communities between OTC and control moss communities for two moss species, Polytrichastrum alpinum and Sanionia uncinata, that make up over 65% of the terrestrial vegetative cover in the area. I found distinct reproductive shifts in P. alpinum under passive warming compared to controls. Moss communities under warming also had substantially larger total invertebrate communities than those in control moss communities, and invertebrate communities were significantly affected by moss species and moss reproductive effort. Further, substantial species-specific thermal differences among contiguous patches of these dominant moss species were revealed. These results suggest that continued warming will differentially impact the reproductive output of Antarctic moss species and is likely to dramatically alter terrestrial ecosystems dynamics from the bottom up. This combined work provides a diverse contribution to the field of epiphyte ecology and biology by providing new insights on how human impacts will affect epiphyte lichen and moss communities across diverse ecosystems, in light of a rapidly changing planet. Title: Examination of Human Impacts on the Biodiversity and Ecology of Lichen and Moss Communities Author: Prather, Hannah Marie Year: 2017 Globally, more than half of the world's population is living in urban areas and it is well accepted that human activities (e.g. climate warming, pollution, landscape homogenization) pose a multitude of threats to ecosystems. Largely, human-related impacts on biodiversity will hold consequences for larger ecological processes and research looking into human impacts on sensitive epiphytic lichen and moss communities is an emerging area of research. While seemingly small, lichen and moss communities exist on nearly every terrestrial ecosystem on Earth and contribute to whole-system processes (e.g. hydrology, mineral cycling, food web energetics) worldwide. To further examine human impacts on epiphytic communities, I conducted three studies examining urbanization and climate warming effects on epiphytic lichen and moss biodiversity and ecology. In the first study I revisited a historic urban lichen community study to assess how urban lichen communities have responded to regional air quality changes occurring over the last nearly two decades. I further investigated, for the first time, the biodiversity of urban tree canopy-dwelling lichen communities in a native coniferous tree species, Pseudotsuga menziesii. I found that urban parks and forested areas harbor a species rich community of lichens epiphytes. Further, I found evidence for the distinct homogenization of urban epiphytic lichen communities, suggesting that expanding beyond simplistic measures of biodiversity to consider community composition and functional biodiversity may be necessary when assessing the ecology and potential ecosystem services of epiphyte communites within urbanizing landscapes. Next, I present the first tall tree canopy study across a regional gradient of urbanization near Portland, Oregon, USA. I found that tall tree canopy epiphyte communities change dramatically along gradients of increasing urbanization, most notably by the transitioning of species functional groups from sensitive, oligotrophic species to a dominance of urban-tolerant, eutrophic species. The implications these dramatic shifts in species composition have on essential PNW ecosystem processes, like N-fixation and canopy microclimate regulation, is still not well understood and is difficult to formally evaluate. However, I find strong evidence that native conifer trees in urban areas may provide a diversity of essential ecosystem services, including providing stratified habitat for epiphyte communities and their associated micro arthropod communities and the scavenging of atmospherically deposited nutrients. Future work is needed to understand how losses in canopy N fixation and species with large biomass (both lichens and bryophytes) will affect nutrient and hydrologic cycling in the PNW region, which continue to undergo rapid growth and urbanization. The final chapter investigates the impacts of passive warming by Open Top Chambers (OTCs) in moss-dominated ecosystems located on the Western Antarctic Peninsula, an area of increasing climate warming. I compared species-specific temperature effects, moss canopy morphology, sexual reproductive effort and invertebrate communities between OTC and control moss communities for two moss species, Polytrichastrum alpinum and Sanionia uncinata, that make up over 65% of the terrestrial vegetative cover in the area. I found distinct reproductive shifts in P. alpinum under passive warming compared to controls. Moss communities under warming also had substantially larger total invertebrate communities than those in control moss communities, and invertebrate communities were significantly affected by moss species and moss reproductive effort. Further, substantial species-specific thermal differences among contiguous patches of these dominant moss species were revealed. These results suggest that continued warming will differentially impact the reproductive output of Antarctic moss species and is likely to dramatically alter terrestrial ecosystems dynamics from the bottom up. This combined work provides a diverse contribution to the field of epiphyte ecology and biology by providing new insights on how human impacts will affect epiphyte lichen and moss communities across diverse ecosystems, in light of a rapidly changing planet. Close

• 283. [Article] Responses of Aquatic Non-Native Species to Novel Predator Cues and Increased Mortality

  Lethal biotic interactions strongly influence the potential for aquatic non-native species to establish and endure in habitats to which they are introduced. Predators in the recipient area, including native ...

  Citation × Citation Citation Abstract Copy Title: Responses of Aquatic Non-Native Species to Novel Predator Cues and Increased Mortality Author: Turner, Brian Christopher Year: 2017 Lethal biotic interactions strongly influence the potential for aquatic non-native species to establish and endure in habitats to which they are introduced. Predators in the recipient area, including native and previously established non-native predators, can prevent establishment, limit habitat use, and reduce abundance of non-native species. Management efforts by humans using methods designed to cause mass mortality (e.g., trapping, biocide applications) can reduce or eradicate non-native populations. However, the impacts of predator and human induced mortality may be mitigated by the behavior or population-level responses of a given non-native species. My dissertation examined the responses of non-native aquatic species to the risk of predation by novel (i.e., no previous exposure) predators in the recipient community and indicators of potential compensatory responses by non-native populations to increased mortality resulting from removal efforts. My dissertation addresses four primary questions. 1) Can first generation, naïve invaders recognize and defend against predators found within the region of invasion through the expression of inducible defenses? 2) Can the overcompensatory potential of a population be predicted though examinations of intraspecific interactions of individuals from the population? 3) What is the relationship between removal effort outcome (i.e., successful or unsuccessful reduction of the target population) and compensatory population responses? 4) Is there a relationship between characteristics of removal efforts that are typically available to managers (e.g., target area size, target area connectivity, removal methodology) and compensatory population responses that could indicate the relative likelihood of compensation resulting from removal efforts? An invading species should be more likely to establish if it can successfully identify and defend against predators in the recipient range, such as through the expression of inducible defenses. Inducible defenses are behavioral or physiological changes that reduce an organism's susceptibility to predation. Through a series of laboratory experiments, I tested whether inducible defenses, in the form of increased burrowing depth, may have benefited the early stage of invasion of Nuttallia obscurata (purple varnish clam), an established Northeast Pacific invader. Specimens of N. obscurata were collected from introduced populations in the Northeast Pacific and from a native population in Japan. The clams were exposed to chemical and physical cues from Northeast Pacific crab predators, including the native Metacarcinus magister (Dungeness crab), an abundant and frequent predator of N. obscurata. While introduced N. obscurata increased their burrowing depth in the physical presence of M. magister, clams collected from their native range showed no such response. This lack of increased burrowing depth by naïve clams in response to a predator native to the newly invaded range, but a significant increase in depth for clams from populations established in the range suggests that while inducible defenses likely did not contribute to the initial establishment of N. obscurata in the Northeast Pacific, they may contribute to their continued persistence and expansion in their introduced range. Some efforts to reduce invasive populations have paradoxically led to population increases. This phenomenon, referred to as overcompensation, occurs when strong negative density-dependent interactions are reduced through increased mortality within a population, resulting in an increase in the population's recruitment rate sufficient to increase the population's overall abundance. Increases in a population's recruitment rate can result from reduced cannibalism of juveniles resulting in lower mortality of new recruits, from increased adult reproductive output, which increases the number of potential recruits, or from reductions in size and/or age at maturity of the unharvested population, which increases the number of reproductive individuals. I predicted the overcompensatory potential of a population of Carcinus maenas (European green crab) in Bodega Harbor, California, using a series of laboratory and field experiments examining intraspecific pressures of adults on juveniles in the population. This measure of intraspecific pressure was used to predict the overcompensatory potential of the population in response to increased mortality from ongoing removal efforts. This prediction was then assessed using pre- and post-removal surveys of juvenile recruitment in Bodega Harbor compared to nearby populations, testing for evidence of overcompensation. While adult C. maenas in Bodega Harbor had limited negative impacts on juveniles, I concluded it was unlikely to result in overcompensation. Relative juvenile abundance did not statistically increase in removal compared to reference populations, consistent with my conclusion from the experiments. Increases in recruitment rates can occur as a result of efforts to remove non-native species. This increase in recruitment can result in overcompensation, but more commonly results in compensation, where recruitment rates increase relative to pre-removal recruitment but does not result in in the population's abundance exceeding pre-removal levels. However, a detailed and accurate prediction of the response of a population to harvest is time consuming and data intensive. This is not feasible for most efforts to eradicate non-native species, which have the greatest chance of success when enacted rapidly after detection. For my final chapter, I performed a literature review and accompanying statistical analysis to determine if typically available information related to the removal effort (site size, site connectivity, and removal technique) could be used to determine increased risk of compensation for a given effort to remove aquatic invasive species. Compensation was closely linked to unsuccessful removal efforts and was observed only among efforts utilizing physical removal methods. However, the frequency with which compensation occurred varied with the exact technique employed, occurring most frequently in removal utilizing electrofishing. Additionally, evidence of compensation was more frequent among larger removal areas with variable connectivity. While other predictors (temperature, effort, etc) might add to the predicative power, the findings of the review provide criteria for managers to determine the relative risk of compensation prior to the start of removal. Further understanding of how invasive species respond to lethal biotic interactions, including anthropogenically mediated control measures, can aid in assessing the risk of invasion for a given species and inform managers of the risk of complications resulting from removal efforts. While inducible defenses may contribute to the long-term success of an introduced species in their recipient range, my findings did not support the idea that inducible defenses triggered by predator cues contributed to their initial introduction in this case. However, research on other non-native species and offspring of previously naïve prey would allow for a clearer picture of the role of inducible defenses in the invasion process. Compensation resulting from removal efforts does not guarantee failure, and certain characteristics of removal efforts seem to indicate increased risk of compensation. Together these components help identify how biotic interactions surrounding mortality risk of an invading species help shape the trajectory of invasion. Title: Responses of Aquatic Non-Native Species to Novel Predator Cues and Increased Mortality Author: Turner, Brian Christopher Year: 2017 Lethal biotic interactions strongly influence the potential for aquatic non-native species to establish and endure in habitats to which they are introduced. Predators in the recipient area, including native and previously established non-native predators, can prevent establishment, limit habitat use, and reduce abundance of non-native species. Management efforts by humans using methods designed to cause mass mortality (e.g., trapping, biocide applications) can reduce or eradicate non-native populations. However, the impacts of predator and human induced mortality may be mitigated by the behavior or population-level responses of a given non-native species. My dissertation examined the responses of non-native aquatic species to the risk of predation by novel (i.e., no previous exposure) predators in the recipient community and indicators of potential compensatory responses by non-native populations to increased mortality resulting from removal efforts. My dissertation addresses four primary questions. 1) Can first generation, naïve invaders recognize and defend against predators found within the region of invasion through the expression of inducible defenses? 2) Can the overcompensatory potential of a population be predicted though examinations of intraspecific interactions of individuals from the population? 3) What is the relationship between removal effort outcome (i.e., successful or unsuccessful reduction of the target population) and compensatory population responses? 4) Is there a relationship between characteristics of removal efforts that are typically available to managers (e.g., target area size, target area connectivity, removal methodology) and compensatory population responses that could indicate the relative likelihood of compensation resulting from removal efforts? An invading species should be more likely to establish if it can successfully identify and defend against predators in the recipient range, such as through the expression of inducible defenses. Inducible defenses are behavioral or physiological changes that reduce an organism's susceptibility to predation. Through a series of laboratory experiments, I tested whether inducible defenses, in the form of increased burrowing depth, may have benefited the early stage of invasion of Nuttallia obscurata (purple varnish clam), an established Northeast Pacific invader. Specimens of N. obscurata were collected from introduced populations in the Northeast Pacific and from a native population in Japan. The clams were exposed to chemical and physical cues from Northeast Pacific crab predators, including the native Metacarcinus magister (Dungeness crab), an abundant and frequent predator of N. obscurata. While introduced N. obscurata increased their burrowing depth in the physical presence of M. magister, clams collected from their native range showed no such response. This lack of increased burrowing depth by naïve clams in response to a predator native to the newly invaded range, but a significant increase in depth for clams from populations established in the range suggests that while inducible defenses likely did not contribute to the initial establishment of N. obscurata in the Northeast Pacific, they may contribute to their continued persistence and expansion in their introduced range. Some efforts to reduce invasive populations have paradoxically led to population increases. This phenomenon, referred to as overcompensation, occurs when strong negative density-dependent interactions are reduced through increased mortality within a population, resulting in an increase in the population's recruitment rate sufficient to increase the population's overall abundance. Increases in a population's recruitment rate can result from reduced cannibalism of juveniles resulting in lower mortality of new recruits, from increased adult reproductive output, which increases the number of potential recruits, or from reductions in size and/or age at maturity of the unharvested population, which increases the number of reproductive individuals. I predicted the overcompensatory potential of a population of Carcinus maenas (European green crab) in Bodega Harbor, California, using a series of laboratory and field experiments examining intraspecific pressures of adults on juveniles in the population. This measure of intraspecific pressure was used to predict the overcompensatory potential of the population in response to increased mortality from ongoing removal efforts. This prediction was then assessed using pre- and post-removal surveys of juvenile recruitment in Bodega Harbor compared to nearby populations, testing for evidence of overcompensation. While adult C. maenas in Bodega Harbor had limited negative impacts on juveniles, I concluded it was unlikely to result in overcompensation. Relative juvenile abundance did not statistically increase in removal compared to reference populations, consistent with my conclusion from the experiments. Increases in recruitment rates can occur as a result of efforts to remove non-native species. This increase in recruitment can result in overcompensation, but more commonly results in compensation, where recruitment rates increase relative to pre-removal recruitment but does not result in in the population's abundance exceeding pre-removal levels. However, a detailed and accurate prediction of the response of a population to harvest is time consuming and data intensive. This is not feasible for most efforts to eradicate non-native species, which have the greatest chance of success when enacted rapidly after detection. For my final chapter, I performed a literature review and accompanying statistical analysis to determine if typically available information related to the removal effort (site size, site connectivity, and removal technique) could be used to determine increased risk of compensation for a given effort to remove aquatic invasive species. Compensation was closely linked to unsuccessful removal efforts and was observed only among efforts utilizing physical removal methods. However, the frequency with which compensation occurred varied with the exact technique employed, occurring most frequently in removal utilizing electrofishing. Additionally, evidence of compensation was more frequent among larger removal areas with variable connectivity. While other predictors (temperature, effort, etc) might add to the predicative power, the findings of the review provide criteria for managers to determine the relative risk of compensation prior to the start of removal. Further understanding of how invasive species respond to lethal biotic interactions, including anthropogenically mediated control measures, can aid in assessing the risk of invasion for a given species and inform managers of the risk of complications resulting from removal efforts. While inducible defenses may contribute to the long-term success of an introduced species in their recipient range, my findings did not support the idea that inducible defenses triggered by predator cues contributed to their initial introduction in this case. However, research on other non-native species and offspring of previously naïve prey would allow for a clearer picture of the role of inducible defenses in the invasion process. Compensation resulting from removal efforts does not guarantee failure, and certain characteristics of removal efforts seem to indicate increased risk of compensation. Together these components help identify how biotic interactions surrounding mortality risk of an invading species help shape the trajectory of invasion. Close