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• 1. [Article] Anthropogenic Effects on the Fouling Community: Impacts of Biological Invasions and Anthropogenic Structures on Community Structure

  Coastal anthropogenic infrastructure has significantly modified nearshore environments. Because these structures often have a strong association with shipping as would be found in ports and harbors, they ...

  Citation × Citation Citation Abstract Copy Title: Anthropogenic Effects on the Fouling Community: Impacts of Biological Invasions and Anthropogenic Structures on Community Structure Author: McClees, Whitney Elizabeth Year: 2017 Coastal anthropogenic infrastructure has significantly modified nearshore environments. Because these structures often have a strong association with shipping as would be found in ports and harbors, they have been identified as invasion hotspots. Due to propagule pressure from shipping and recreational boating and suitable uncolonized substrate that provides a refuge from native predators, a greater number of non-native species have been found on these structures compared to nearby natural substrate. The mechanisms that limit the spread of non-native species from anthropogenic structures to natural substrate have been explored for several taxa at a species-specific level, but less so from an overall community perspective. Predation has been identified as one of the biotic interactions limiting invasion success. In addition to predation, dispersal ability may also prevent the spread of non-native species from anthropogenic structures to natural substrate. This thesis addresses how these two mechanisms interact to limit the spread of non-native species from anthropogenic structures to natural substrate and how that alters overall community composition. I aimed to explore differences between communities inside and outside of a marina and determine the extent to which predator and dispersal limitation were structuring these communities. I used a three-factor design, deploying seven unglazed ceramic tiles per each treatment combination of 1) in versus outside a marina in Yaquina Bay, Oregon; 2) cage keeping out predators greater than the mesh size, no cage, or partial cage; 3) fixed near the substrata (benthic) versus suspended 1 meter below the surface. I also transplanted caged, suspended tiles of either adults or recruits from inside the marina to benthic and suspended caging treatments outside of the marina. These tiles allowed me to examine predation when dispersal limitation was not a factor for the community inside the marina, i.e. what happens to both recruits and adults if they can get outside of the marina. I found that the communities inside and outside of the marina were different and the data suggest that both predation and dispersal limitation interact to limit the spread of non-native species. Additionally, I found that mesopredators that could fit through the caging may be influencing predation results and community structure. This research addresses gaps in scientific knowledge regarding the mechanisms that prevent or facilitate the spread of non-native species. Future work could include the further exploration of mesopredation as an important factor in limiting the spread of non-native species and exploring dispersal limitation more in depth as well as broadening the geographic scope to see if the same trends hold true across bays and bioregions. Title: Anthropogenic Effects on the Fouling Community: Impacts of Biological Invasions and Anthropogenic Structures on Community Structure Author: McClees, Whitney Elizabeth Year: 2017 Coastal anthropogenic infrastructure has significantly modified nearshore environments. Because these structures often have a strong association with shipping as would be found in ports and harbors, they have been identified as invasion hotspots. Due to propagule pressure from shipping and recreational boating and suitable uncolonized substrate that provides a refuge from native predators, a greater number of non-native species have been found on these structures compared to nearby natural substrate. The mechanisms that limit the spread of non-native species from anthropogenic structures to natural substrate have been explored for several taxa at a species-specific level, but less so from an overall community perspective. Predation has been identified as one of the biotic interactions limiting invasion success. In addition to predation, dispersal ability may also prevent the spread of non-native species from anthropogenic structures to natural substrate. This thesis addresses how these two mechanisms interact to limit the spread of non-native species from anthropogenic structures to natural substrate and how that alters overall community composition. I aimed to explore differences between communities inside and outside of a marina and determine the extent to which predator and dispersal limitation were structuring these communities. I used a three-factor design, deploying seven unglazed ceramic tiles per each treatment combination of 1) in versus outside a marina in Yaquina Bay, Oregon; 2) cage keeping out predators greater than the mesh size, no cage, or partial cage; 3) fixed near the substrata (benthic) versus suspended 1 meter below the surface. I also transplanted caged, suspended tiles of either adults or recruits from inside the marina to benthic and suspended caging treatments outside of the marina. These tiles allowed me to examine predation when dispersal limitation was not a factor for the community inside the marina, i.e. what happens to both recruits and adults if they can get outside of the marina. I found that the communities inside and outside of the marina were different and the data suggest that both predation and dispersal limitation interact to limit the spread of non-native species. Additionally, I found that mesopredators that could fit through the caging may be influencing predation results and community structure. This research addresses gaps in scientific knowledge regarding the mechanisms that prevent or facilitate the spread of non-native species. Future work could include the further exploration of mesopredation as an important factor in limiting the spread of non-native species and exploring dispersal limitation more in depth as well as broadening the geographic scope to see if the same trends hold true across bays and bioregions. Close

• 2. [Article] The Limitation of Spread of Non-native Marine Invertebrates from Artificial Structures to Natural Habitats

  Artificial structures created for aquatic anthropogenic activities are often colonized and fouled by many non-native species, few of which have invaded natural areas. Some research has indicated predation ...

  Citation × Citation Citation Abstract Copy Title: The Limitation of Spread of Non-native Marine Invertebrates from Artificial Structures to Natural Habitats Author: McClees, Whitney, de Rivera, Catherine E. Year: 2017 Artificial structures created for aquatic anthropogenic activities are often colonized and fouled by many non-native species, few of which have invaded natural areas. Some research has indicated predation is limiting the spread of non-native species, while other research has indicated that it is dispersal-driven. To determine the effects of dispersal limitation and predation on the risk to surrounding habitats of being invaded by non-native species established on marina docks, we used a three-factor design, deploying seven unglazed ceramic panels per each treatment combination of 1) inside versus outside a marina in Yaquina Bay, Oregon; 2) cage keeping out predators >mesh size, no cage, or partial cage; 3) fixed near the substrata (benthic) versus floating 1m below the surface. We also moved panels of either adults or recruits from floating in the marina to any of the treatment combinations to test if they would have lower survival outside by the substrate. The community on panels suspended in the marina only minimally overlapped with panels outside of the marina near the bottom. However, there was no difference between the cage treatments, which suggests that predators excluded by the cages are not responsible for the observed differences in species richness or composition. Title: The Limitation of Spread of Non-native Marine Invertebrates from Artificial Structures to Natural Habitats Author: McClees, Whitney, de Rivera, Catherine E. Year: 2017 Artificial structures created for aquatic anthropogenic activities are often colonized and fouled by many non-native species, few of which have invaded natural areas. Some research has indicated predation is limiting the spread of non-native species, while other research has indicated that it is dispersal-driven. To determine the effects of dispersal limitation and predation on the risk to surrounding habitats of being invaded by non-native species established on marina docks, we used a three-factor design, deploying seven unglazed ceramic panels per each treatment combination of 1) inside versus outside a marina in Yaquina Bay, Oregon; 2) cage keeping out predators >mesh size, no cage, or partial cage; 3) fixed near the substrata (benthic) versus floating 1m below the surface. We also moved panels of either adults or recruits from floating in the marina to any of the treatment combinations to test if they would have lower survival outside by the substrate. The community on panels suspended in the marina only minimally overlapped with panels outside of the marina near the bottom. However, there was no difference between the cage treatments, which suggests that predators excluded by the cages are not responsible for the observed differences in species richness or composition. Close

• 3. [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

• 4. [Article] Biofouling Management in the Pacific Northwest and Predation on Native versus Non-native Ascidians

  Marine non-native species threaten economic and environmental health, making it crucial to understand factors that make them successful. Research on these species, therefore, allows for greater preparedness ...

  Citation × Citation Citation Abstract Copy Title: Biofouling Management in the Pacific Northwest and Predation on Native versus Non-native Ascidians Author: Kincaid, Erin Suzanne Year: 2016 Marine non-native species threaten economic and environmental health, making it crucial to understand factors that make them successful. Research on these species, therefore, allows for greater preparedness and informed management of biological invasions and increases understanding of elements structuring biological communities. Among the marine non-native species, and particularly the fouling community, non-native ascidians are a taxon of particular concern because they can crowd out native benthic species and smother mariculture products. This thesis addresses management for ascidians and other fouling organisms and includes research on the invasiveness of this taxon in addition to the invasibility of recipient fouling communities. On the West Coast of the U.S., limited efforts have been made to coordinate biofouling management across states, despite the myriad vectors increasing propagule pressure over time along coastal states. Building on recent state and local efforts, I developed a Pacific Regional Biofouling Plan for the states of Oregon and Washington to help start a consensus-driven process by which these states could create a forum for more comprehensive coordination efforts, following California's lead. As states address authority gaps, the biofouling management framework I've written is meant to be used to guide the conversation between managers as various stages of coastal management are realized. To better inform the scope and efficacy of management and regulatory efforts, the study of invasions ecology asks and aims to answer questions regarding recipient community interactions and characteristics of the non-native species themselves. Studies that identify characteristics that make ascidians successful (invasiveness) and determine the influence native communities have on their success (invasibility) are important for assessing overall risk of establishment and spread from non-native ascidians. Therefore, I aimed to: 1) explore the hypothesis that fouling communities on suspended, artificial structures are more invasible than benthic habitats; and 2) identify characteristics influencing predation patterns on the native Distaplia occidentalis versus non-native ascidian species using mensurative and experimental studies in Charleston Marina, Oregon. I conducted a series of feeding assays, surveys, and a caloric content analysis. Feeding assays were conducted with a suite of predators. The flatworm predator (Eurylepta leoparda) was found to be highly selective on the native ascidian Distaplia occidentalis, and only preyed on whole colony samples. Feeding assay data suggest that test (tunic) structure or thickness may be an influential factor affecting nudibranch (Hermissenda crassicornis) predation rates on native versus non-native ascidians, with greater predation on the native ascidian species. Non-native ascidians may escape predation in floating but not benthic environments on the Oregon coast due to their palatability characteristics, likely tunic structure and low caloric content. In this case, this suite of predators may indirectly facilitate the invasion of docks but provide at least partial resistance to the invasion of natural benthic areas. The chapters herein address gaps in management and scientific knowledge regarding non-native species of the marine fouling community. Future work enhanced by my efforts could include the development of the coastal biofouling management plan, coordinated by the Western Regional Panel on Aquatic Invasive Species Coastal Committee, and broadening the geographic and taxonomic scope of my research with a more comprehensive study of predator-prey interactions involving non-native ascidians and a diverse suite of predators. These interactions may be an important factor in explaining the success of ascidians and other fouling organisms on floating structures and lack of success on nearby benthic substrata. Title: Biofouling Management in the Pacific Northwest and Predation on Native versus Non-native Ascidians Author: Kincaid, Erin Suzanne Year: 2016 Marine non-native species threaten economic and environmental health, making it crucial to understand factors that make them successful. Research on these species, therefore, allows for greater preparedness and informed management of biological invasions and increases understanding of elements structuring biological communities. Among the marine non-native species, and particularly the fouling community, non-native ascidians are a taxon of particular concern because they can crowd out native benthic species and smother mariculture products. This thesis addresses management for ascidians and other fouling organisms and includes research on the invasiveness of this taxon in addition to the invasibility of recipient fouling communities. On the West Coast of the U.S., limited efforts have been made to coordinate biofouling management across states, despite the myriad vectors increasing propagule pressure over time along coastal states. Building on recent state and local efforts, I developed a Pacific Regional Biofouling Plan for the states of Oregon and Washington to help start a consensus-driven process by which these states could create a forum for more comprehensive coordination efforts, following California's lead. As states address authority gaps, the biofouling management framework I've written is meant to be used to guide the conversation between managers as various stages of coastal management are realized. To better inform the scope and efficacy of management and regulatory efforts, the study of invasions ecology asks and aims to answer questions regarding recipient community interactions and characteristics of the non-native species themselves. Studies that identify characteristics that make ascidians successful (invasiveness) and determine the influence native communities have on their success (invasibility) are important for assessing overall risk of establishment and spread from non-native ascidians. Therefore, I aimed to: 1) explore the hypothesis that fouling communities on suspended, artificial structures are more invasible than benthic habitats; and 2) identify characteristics influencing predation patterns on the native Distaplia occidentalis versus non-native ascidian species using mensurative and experimental studies in Charleston Marina, Oregon. I conducted a series of feeding assays, surveys, and a caloric content analysis. Feeding assays were conducted with a suite of predators. The flatworm predator (Eurylepta leoparda) was found to be highly selective on the native ascidian Distaplia occidentalis, and only preyed on whole colony samples. Feeding assay data suggest that test (tunic) structure or thickness may be an influential factor affecting nudibranch (Hermissenda crassicornis) predation rates on native versus non-native ascidians, with greater predation on the native ascidian species. Non-native ascidians may escape predation in floating but not benthic environments on the Oregon coast due to their palatability characteristics, likely tunic structure and low caloric content. In this case, this suite of predators may indirectly facilitate the invasion of docks but provide at least partial resistance to the invasion of natural benthic areas. The chapters herein address gaps in management and scientific knowledge regarding non-native species of the marine fouling community. Future work enhanced by my efforts could include the development of the coastal biofouling management plan, coordinated by the Western Regional Panel on Aquatic Invasive Species Coastal Committee, and broadening the geographic and taxonomic scope of my research with a more comprehensive study of predator-prey interactions involving non-native ascidians and a diverse suite of predators. These interactions may be an important factor in explaining the success of ascidians and other fouling organisms on floating structures and lack of success on nearby benthic substrata. Close

• 5. [Article] Fish Species Introductions Provide Novel Insights into the Patterns and Drivers of Phylogenetic Structure in Freshwaters

  Despite long-standing interest of terrestrial ecologists, freshwater ecosystems are a fertile, yet unappreciated, testing ground for applying community phylogenetics to uncover mechanisms of species assembly. ...

  Citation × Citation Citation Abstract Copy Title: Fish Species Introductions Provide Novel Insights into the Patterns and Drivers of Phylogenetic Structure in Freshwaters Author: Strecker, Angela L., Olden, Julian D. Year: 2014 Despite long-standing interest of terrestrial ecologists, freshwater ecosystems are a fertile, yet unappreciated, testing ground for applying community phylogenetics to uncover mechanisms of species assembly. We quantify phylogenetic clustering and overdispersion of native and non-native fishes of a large river basin in the American Southwest to test for the mechanisms (environmental filtering versus competitive exclusion) and spatial scales influencing community structure. Contrary to expectations, non-native species were phylogenetically clustered and related to natural environmental conditions, whereas native species were not phylogenetically structured, likely reflecting human-related changes to the basin. The species that are most invasive (in terms of ecological impacts) tended to be the most phylogenetically divergent from natives across watersheds, but not within watersheds, supporting the hypothesis that Darwin's naturalization conundrum is driven by the spatial scale. Phylogenetic distinctiveness may facilitate non-native establishment at regional scales, but environmental filtering restricts local membership to closely related species with physiological tolerances for current environments. By contrast, native species may have been phylogenetically clustered in historical times, but species loss from contemporary populations by anthropogenic activities has likely shaped the phylogenetic signal. Our study implies that fundamental mechanisms of community assembly have changed, with fundamental consequences for the biogeography of both native and non-native species. Title: Fish Species Introductions Provide Novel Insights into the Patterns and Drivers of Phylogenetic Structure in Freshwaters Author: Strecker, Angela L., Olden, Julian D. Year: 2014 Despite long-standing interest of terrestrial ecologists, freshwater ecosystems are a fertile, yet unappreciated, testing ground for applying community phylogenetics to uncover mechanisms of species assembly. We quantify phylogenetic clustering and overdispersion of native and non-native fishes of a large river basin in the American Southwest to test for the mechanisms (environmental filtering versus competitive exclusion) and spatial scales influencing community structure. Contrary to expectations, non-native species were phylogenetically clustered and related to natural environmental conditions, whereas native species were not phylogenetically structured, likely reflecting human-related changes to the basin. The species that are most invasive (in terms of ecological impacts) tended to be the most phylogenetically divergent from natives across watersheds, but not within watersheds, supporting the hypothesis that Darwin's naturalization conundrum is driven by the spatial scale. Phylogenetic distinctiveness may facilitate non-native establishment at regional scales, but environmental filtering restricts local membership to closely related species with physiological tolerances for current environments. By contrast, native species may have been phylogenetically clustered in historical times, but species loss from contemporary populations by anthropogenic activities has likely shaped the phylogenetic signal. Our study implies that fundamental mechanisms of community assembly have changed, with fundamental consequences for the biogeography of both native and non-native species. Close

• 6. [Article] Non-native Mangroves of Moloka’i, Hawai’i: A Socio-ecological Analysis

  Invasive species are recognized as a leading threat to biodiversity and their management is expensive, time consuming, and labor intensive. Therefore, it is important to review both benefits and detriments ...

  Citation × Citation Citation Abstract Copy Title: Non-native Mangroves of Moloka’i, Hawai’i: A Socio-ecological Analysis Author: Lewis, Casey Year: 2017 Invasive species are recognized as a leading threat to biodiversity and their management is expensive, time consuming, and labor intensive. Therefore, it is important to review both benefits and detriments of the species to inform appropriate management decisions. Red mangrove was introduced to Moloka'i, Hawaii in 1902 to mitigate the effects of soil erosion and has since spread along the coast and to adjacent islands creating novel habitat. This study assessed both biological services and social attitudes towards Moloka'i's non-native mangroves to provide a comprehensive evaluation of the species. Zooplankton community structure was examined in mangrove and non-mangrove sites and found to be different depending on the amount of mangroves present. Surveys of Moloka'i residents revealed no majority attitude towards mangroves nor support for complete eradication but residents were highly supportive of active management. Approaching other novel ecosystems created by established non-native species from a socio-ecological view may become more crucial as changes in climate, demographics, and environmental conditions further complicate effects. In our globalized world, non-native species introductions will continue to have both positive and negative effects that vary over time. Therefore, assessment of both social and ecological perspectives may lead to more effective management with greater community support. Title: Non-native Mangroves of Moloka’i, Hawai’i: A Socio-ecological Analysis Author: Lewis, Casey Year: 2017 Invasive species are recognized as a leading threat to biodiversity and their management is expensive, time consuming, and labor intensive. Therefore, it is important to review both benefits and detriments of the species to inform appropriate management decisions. Red mangrove was introduced to Moloka'i, Hawaii in 1902 to mitigate the effects of soil erosion and has since spread along the coast and to adjacent islands creating novel habitat. This study assessed both biological services and social attitudes towards Moloka'i's non-native mangroves to provide a comprehensive evaluation of the species. Zooplankton community structure was examined in mangrove and non-mangrove sites and found to be different depending on the amount of mangroves present. Surveys of Moloka'i residents revealed no majority attitude towards mangroves nor support for complete eradication but residents were highly supportive of active management. Approaching other novel ecosystems created by established non-native species from a socio-ecological view may become more crucial as changes in climate, demographics, and environmental conditions further complicate effects. In our globalized world, non-native species introductions will continue to have both positive and negative effects that vary over time. Therefore, assessment of both social and ecological perspectives may lead to more effective management with greater community support. Close

• 7. [Article] Non-native Mangroves of Moloka'i, Hawai'i: A Socio-ecological Analysis

  Invasive species are recognized as a leading threat to ecosystems and their management is expensive, time consuming, and labor intensive. Therefore, it is important to review both benefits and detriments ...

  Citation × Citation Citation Abstract Copy Title: Non-native Mangroves of Moloka'i, Hawai'i: A Socio-ecological Analysis Author: Lewis, Casey Year: 2017 Invasive species are recognized as a leading threat to ecosystems and their management is expensive, time consuming, and labor intensive. Therefore, it is important to review both benefits and detriments of the species to inform appropriate management decisions. Red mangrove was introduced to Moloka'i, Hawaii in 1902 to mitigate the effects of soil erosion and has since spread along the coast and to adjacent islands creating novel habitat. This study assessed both biological services and social attitudes towards Moloka'i's non-native mangroves to provide a comprehensive evaluation of the species. Zooplankton community structure was examined in mangrove and non-mangrove sites and found to be different depending on the amount of mangroves present. Surveys of Moloka'i residents revealed no majority attitude towards mangroves nor support for complete eradication but residents were highly supportive of active management. Approaching other novel ecosystems created by established non-native species from a socio-ecological view may become more crucial as changes in climate, demographics, and environmental conditions further complicate effects. In our globalized world, non-native species introductions will continue to have both positive and negative effects that vary over time. Therefore, assessment of both social and ecological perspectives may lead to more effective management with greater community support. Title: Non-native Mangroves of Moloka'i, Hawai'i: A Socio-ecological Analysis Author: Lewis, Casey Year: 2017 Invasive species are recognized as a leading threat to ecosystems and their management is expensive, time consuming, and labor intensive. Therefore, it is important to review both benefits and detriments of the species to inform appropriate management decisions. Red mangrove was introduced to Moloka'i, Hawaii in 1902 to mitigate the effects of soil erosion and has since spread along the coast and to adjacent islands creating novel habitat. This study assessed both biological services and social attitudes towards Moloka'i's non-native mangroves to provide a comprehensive evaluation of the species. Zooplankton community structure was examined in mangrove and non-mangrove sites and found to be different depending on the amount of mangroves present. Surveys of Moloka'i residents revealed no majority attitude towards mangroves nor support for complete eradication but residents were highly supportive of active management. Approaching other novel ecosystems created by established non-native species from a socio-ecological view may become more crucial as changes in climate, demographics, and environmental conditions further complicate effects. In our globalized world, non-native species introductions will continue to have both positive and negative effects that vary over time. Therefore, assessment of both social and ecological perspectives may lead to more effective management with greater community support. Close

• 8. [Article] A Four Year Study of the Effects of Substrate Depth on the Survival of Different Plant Species in Portland, OR

  The changes in green roof plant species composition was observed after a four year time period. Twelve plots were planted in either native or non-native species at two different substrate depths according ...

  Citation × Citation Citation Abstract Copy Title: A Four Year Study of the Effects of Substrate Depth on the Survival of Different Plant Species in Portland, OR Author: Viray, Arjun Year: 2017 The changes in green roof plant species composition was observed after a four year time period. Twelve plots were planted in either native or non-native species at two different substrate depths according to a 2 by 2 factorial design. The list of original species from installation was used to identify which species survived over time in their respective plots. After four years, 9 out of 12 native species and 13 out of 14 species survived. The high survival of 26 original species must be compared to the colonization of 68 new species. With minimal maintenance, new species colonized plots. Extensive plots were composed of 33% original planting and 67% colonizing species. Intensive plots were composed of 24% original plantings and 76% new species. The original species that had the highest survival were Sedums and Festuca. Authors: Arjun Vijay, Olyssa Starry Presenter: Arjun Viray Title: A Four Year Study of the Effects of Substrate Depth on the Survival of Different Plant Species in Portland, OR Author: Viray, Arjun Year: 2017 The changes in green roof plant species composition was observed after a four year time period. Twelve plots were planted in either native or non-native species at two different substrate depths according to a 2 by 2 factorial design. The list of original species from installation was used to identify which species survived over time in their respective plots. After four years, 9 out of 12 native species and 13 out of 14 species survived. The high survival of 26 original species must be compared to the colonization of 68 new species. With minimal maintenance, new species colonized plots. Extensive plots were composed of 33% original planting and 67% colonizing species. Intensive plots were composed of 24% original plantings and 76% new species. The original species that had the highest survival were Sedums and Festuca. Authors: Arjun Vijay, Olyssa Starry Presenter: Arjun Viray Close

• 9. [Article] Ecosystem Recovery in Estuarine Wetlands of the Columbia River Estuary

  In the restoration of tidal wetland ecosystems, potential drivers of plant community development range from biotic controls (e.g. plant competition, seed dispersal) to abiotic controls (e.g. tidal flooding, ...

  Citation × Citation Citation Abstract Copy Title: Ecosystem Recovery in Estuarine Wetlands of the Columbia River Estuary Author: Kidd, Sarah Ann Year: 2017 In the restoration of tidal wetland ecosystems, potential drivers of plant community development range from biotic controls (e.g. plant competition, seed dispersal) to abiotic controls (e.g. tidal flooding, salinity levels). How these controls influence the success of tidal wetland restoration are only partly understood, but have important implications for wetland habitat recovery. Specifically, the extent to which the existing native and non-native seed banks in tidally reconnected wetlands interact with these controls is not clear, yet the potential success of passive restoration methods depends upon this understanding. For a 54-year chronosequence of eleven tidal wetland restoration sites in the Lower Columbia River of western Oregon, USA, it was hypothesized that native plant species and soil properties would show trends approaching reference levels within 3 to 20 years post-restoration and that lower elevation wetland areas within restored sites would exhibit a greater native species abundance and similarity to reference sites, compared with restored high elevation wetland areas. Results indicated that plant species richness, soil organic matter, bulk density, pH, and salinity conditions among the restoration sites reached reference wetland ranges within 3-6 years post-tidal reconnection. The mid-low marsh elevation zones (2.5 m) remained dominated by non-native species Phalaris arundinacea and Juncus effusus subsp. effusus. To investigate the mechanisms driving these non-native plant invasions, it was hypothesized that native and non-native wetland plant community distributions would be reflective both of their abundance in the seed bank and of their germination tolerance to wetland tidal flooding and salinity conditions. Using a factorial study design of three tidal conditions by three salinity levels, these hypotheses were tested in the greenhouse. Overall, non-native seeds were found to significantly outnumber native seeds in both seed banks. In the greenhouse, P. arundinacea and J. effusus were found to germinate more readily out of the seed bank under freshwater high-marsh flooding (1 hour a day) treatments as compared to oligohaline (3 ppt) mid-low marsh flooding (3-6 hours twice a day) treatments and to brackish salinity (10 ppt) treatments. Dominant native wetland species, Carex lyngbyei and Schoenoplectus lacustris, germination were not found to vary significantly among the treatments (p > 0.10). These results indicate that the salinity and flooding gradients within these restored marshes suppress germination of the non-native species in the low-mid marsh but not in the high marsh, where they are likely able to outcompete the native species due to their dominance in the seed bank. The implications of these results for passive tidal wetland restoration efforts are that both seed bank composition and species-specific tolerances to restored tidal flooding and salinity gradients are key mechanisms driving native and non-native plant community development and resilience. Title: Ecosystem Recovery in Estuarine Wetlands of the Columbia River Estuary Author: Kidd, Sarah Ann Year: 2017 In the restoration of tidal wetland ecosystems, potential drivers of plant community development range from biotic controls (e.g. plant competition, seed dispersal) to abiotic controls (e.g. tidal flooding, salinity levels). How these controls influence the success of tidal wetland restoration are only partly understood, but have important implications for wetland habitat recovery. Specifically, the extent to which the existing native and non-native seed banks in tidally reconnected wetlands interact with these controls is not clear, yet the potential success of passive restoration methods depends upon this understanding. For a 54-year chronosequence of eleven tidal wetland restoration sites in the Lower Columbia River of western Oregon, USA, it was hypothesized that native plant species and soil properties would show trends approaching reference levels within 3 to 20 years post-restoration and that lower elevation wetland areas within restored sites would exhibit a greater native species abundance and similarity to reference sites, compared with restored high elevation wetland areas. Results indicated that plant species richness, soil organic matter, bulk density, pH, and salinity conditions among the restoration sites reached reference wetland ranges within 3-6 years post-tidal reconnection. The mid-low marsh elevation zones (2.5 m) remained dominated by non-native species Phalaris arundinacea and Juncus effusus subsp. effusus. To investigate the mechanisms driving these non-native plant invasions, it was hypothesized that native and non-native wetland plant community distributions would be reflective both of their abundance in the seed bank and of their germination tolerance to wetland tidal flooding and salinity conditions. Using a factorial study design of three tidal conditions by three salinity levels, these hypotheses were tested in the greenhouse. Overall, non-native seeds were found to significantly outnumber native seeds in both seed banks. In the greenhouse, P. arundinacea and J. effusus were found to germinate more readily out of the seed bank under freshwater high-marsh flooding (1 hour a day) treatments as compared to oligohaline (3 ppt) mid-low marsh flooding (3-6 hours twice a day) treatments and to brackish salinity (10 ppt) treatments. Dominant native wetland species, Carex lyngbyei and Schoenoplectus lacustris, germination were not found to vary significantly among the treatments (p > 0.10). These results indicate that the salinity and flooding gradients within these restored marshes suppress germination of the non-native species in the low-mid marsh but not in the high marsh, where they are likely able to outcompete the native species due to their dominance in the seed bank. The implications of these results for passive tidal wetland restoration efforts are that both seed bank composition and species-specific tolerances to restored tidal flooding and salinity gradients are key mechanisms driving native and non-native plant community development and resilience. Close

• 10. [Article] Increased Habitat Connectivity Homogenizes Freshwater Communities: Historical and Landscape Perspectives

  Increases in habitat connectivity can have consequences for taxonomic, functional, and genetic diversity of communities. Previously isolated aquatic habitats were connected with canals and pipelines in ...

  Citation × Citation Citation Abstract Copy Title: Increased Habitat Connectivity Homogenizes Freshwater Communities: Historical and Landscape Perspectives Author: Strecker, Angela L., Brittain, Jeffrey Thomas Year: 2017 Increases in habitat connectivity can have consequences for taxonomic, functional, and genetic diversity of communities. Previously isolated aquatic habitats were connected with canals and pipelines in the largest water development project in the US history, the Columbia Basin Project (CBP; eastern Washington, USA), which also altered environmental conditions; however, the ecological consequences are largely unknown. Using a historical dataset, we examined long-term patterns in zooplankton communities, water chemistry and clarity, testing the hypothesis that increased connectivity will result in taxonomic homogenization. Further, we tested contemporary drivers of communities using a comprehensive set of environmental and landscape variables. Waterbodies were sampled for zooplankton community composition as well as physical and chemical variables inside and outside the CBP using methods consistent with historical studies. We found significant declines in salinity inside the CBP, whereas changes in water clarity were prevalent across all waterbodies. Increased connectivity via canals homogenized zooplankton communities over time, as well as increasing regional richness. Other long-term changes in zooplankton communities may be related to climate change, invasive species, and land-use changes. Synthesis and applications. Though canals may offer species spatial refugia, homogenization may decrease resilience to environmental stressors. These new hybrid aquatic landscapes, or hydroscapes, should be considered carefully in future water development, including specific plans for monitoring of species and environmental conditions, as well as mitigation of undesirable conditions and/or non-native species. Title: Increased Habitat Connectivity Homogenizes Freshwater Communities: Historical and Landscape Perspectives Author: Strecker, Angela L., Brittain, Jeffrey Thomas Year: 2017 Increases in habitat connectivity can have consequences for taxonomic, functional, and genetic diversity of communities. Previously isolated aquatic habitats were connected with canals and pipelines in the largest water development project in the US history, the Columbia Basin Project (CBP; eastern Washington, USA), which also altered environmental conditions; however, the ecological consequences are largely unknown. Using a historical dataset, we examined long-term patterns in zooplankton communities, water chemistry and clarity, testing the hypothesis that increased connectivity will result in taxonomic homogenization. Further, we tested contemporary drivers of communities using a comprehensive set of environmental and landscape variables. Waterbodies were sampled for zooplankton community composition as well as physical and chemical variables inside and outside the CBP using methods consistent with historical studies. We found significant declines in salinity inside the CBP, whereas changes in water clarity were prevalent across all waterbodies. Increased connectivity via canals homogenized zooplankton communities over time, as well as increasing regional richness. Other long-term changes in zooplankton communities may be related to climate change, invasive species, and land-use changes. Synthesis and applications. Though canals may offer species spatial refugia, homogenization may decrease resilience to environmental stressors. These new hybrid aquatic landscapes, or hydroscapes, should be considered carefully in future water development, including specific plans for monitoring of species and environmental conditions, as well as mitigation of undesirable conditions and/or non-native species. Close