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• 1. [Article] The Biology of Eastern Kingbirds at Malheur National Wildlife Refuge: Survival, Reproduction, and Testosterone Secretion

  This dissertation presents the results of a study that I undertook to better understand the breeding biology of Eastern Kingbirds (hereafter, kingbirds) at Malheur National Wildlife Refuge in southeastern ...

  Citation × Citation Citation Abstract Copy Title: The Biology of Eastern Kingbirds at Malheur National Wildlife Refuge: Survival, Reproduction, and Testosterone Secretion Author: Redmond, Lucas J. Year: 2015 This dissertation presents the results of a study that I undertook to better understand the breeding biology of Eastern Kingbirds (hereafter, kingbirds) at Malheur National Wildlife Refuge in southeastern Oregon from 2003 to 2009. Kingbirds are long-distance migratory songbirds that breed across much of North America. This species is socially monogamous but, via frequent extra-pair copulations, is genetically polygamous. Kingbirds exhibit relatively high breeding site fidelity, often returning to the same tree to nest in subsequent years. Both members of a pair provide parental care, but there are often specific duties performed by both male and female kingbirds. For example, males typically perform much of the vigilant nest defense that this species is well known for and contribute, to an extent, to the feeding of nestlings. Females, on the other hand, are entirely responsible for incubating and brooding. Beginning in 2002, most adults within the population of kingbirds at Malheur had been banded with a unique combination of three colored plastic leg bands and an aluminum USFWS band. Also, as many nests were located in each year as possible, and any young that survived to fledging age were banded with a unique combination of leg bands. Considerable effort was then spent each year to locate as many banded individuals as possible, which allowed me to document adult and juvenile survival. Annual survival rates of adult male and female kingbirds in the population at Malheur did not differ and were relatively high at approximately 0.65. Juvenile survival rate was approximately 0.29, indicating that slightly less than one-third of nestlings survive the interval between fledging and their first potential breeding season. Resighting probability was high for both sexes, although higher for adult males (0.94) than for adult female (0.84). The latter finding is consistent with the higher site fidelity of males than of females. Resighting probabilities for juveniles were much lower at 0.68 and 0.40 for males and females, respectively. Again, this was expected because natal site fidelity is typically much lower than breeding site fidelity. Compared to most other reports, resighting probability and return rates of juvenile kingbirds was high, presumably because the riparian habitats where kingbirds breed at Malheur function as an ecological island surrounded by, for kingbirds, unsuitable high desert habitat. Thus, unless they disperse very long distances, the only option for juvenile kingbirds is to begin nesting on the refuge. The collection of blood samples from birds has become an increasingly common practice in ornithology. The data that can be gained from these samples allow a number of interesting questions to be asked such as understanding the genetic mating system of a species, patterns of hormone secretion, and discerning migratory pathways via stable isotope analysis. The volume of blood collected is usually small and was assumed to cause no long-term negative effects on sampled individuals. However, few studies have rigorously examined the effect of blood sampling on survival. I used a multistate mark-recapture analysis to assess the effect blood sampling on annual survival of kingbirds by combining the annual survival data described above with whether or not individuals were subjected to blood sample collection. The results of this analysis indicated that blood sampling had no effect on annual survival rates of kingbirds. Whether or not this is the case for other species remains to be seen. However, my results support the assumption that when done correctly, blood sampling has little to no long-term negative effects on birds. Comparative analyses of many bird species show that testosterone secretion exhibits fairly predictable patterns among breeding birds of different mating systems. Monogamous species reach a peak during mate attraction and the period of female fertility which is then followed by a sharp decline when young are in the nest. By contrast, males of polygynous species tend to maintain higher levels of testosterone throughout the breeding season to, presumably, maximize opportunities for acquiring additional mates. Kingbirds are socially monogamous but cryptically polygamous. However, because of high rates of extra-pair paternity, variation in reproductive success among males is much higher than what is expected for a monogamous species, and, instead, is similar to what has been reported for polygynous species. Therefore from 2005 to 2009 I measured testosterone concentrations from the blood samples collected from male kingbirds to determine the breeding season profile of testosterone secretion in kingbirds and to understand the factors that influence testosterone variation among individuals. Contrary to expected for a monogamous species, the testosterone profile of kingbirds did not exhibit the brief peak in testosterone followed by a precipitous decline. Testosterone peaked early in the season, but declined very slowly as the nest cycle progressed. I attributed this gradual decline in testosterone to the cryptically polygynous nature of the kingbird extra-pair mating system. I also found substantial variation in testosterone concentration among male kingbirds and was able to identify several factors contributing to this variation. As expected, testosterone declined as the breeding season progressed (independent of stage in nest cycle) and nest density increased, while increases in testosterone were correlated with the number of fertile females within the population. This suggests that male kingbirds were capable of modulating testosterone concentrations to both cope with an increase in conspecific density, but also to ready themselves for times when extra-pair copulations were more likely. Title: The Biology of Eastern Kingbirds at Malheur National Wildlife Refuge: Survival, Reproduction, and Testosterone Secretion Author: Redmond, Lucas J. Year: 2015 This dissertation presents the results of a study that I undertook to better understand the breeding biology of Eastern Kingbirds (hereafter, kingbirds) at Malheur National Wildlife Refuge in southeastern Oregon from 2003 to 2009. Kingbirds are long-distance migratory songbirds that breed across much of North America. This species is socially monogamous but, via frequent extra-pair copulations, is genetically polygamous. Kingbirds exhibit relatively high breeding site fidelity, often returning to the same tree to nest in subsequent years. Both members of a pair provide parental care, but there are often specific duties performed by both male and female kingbirds. For example, males typically perform much of the vigilant nest defense that this species is well known for and contribute, to an extent, to the feeding of nestlings. Females, on the other hand, are entirely responsible for incubating and brooding. Beginning in 2002, most adults within the population of kingbirds at Malheur had been banded with a unique combination of three colored plastic leg bands and an aluminum USFWS band. Also, as many nests were located in each year as possible, and any young that survived to fledging age were banded with a unique combination of leg bands. Considerable effort was then spent each year to locate as many banded individuals as possible, which allowed me to document adult and juvenile survival. Annual survival rates of adult male and female kingbirds in the population at Malheur did not differ and were relatively high at approximately 0.65. Juvenile survival rate was approximately 0.29, indicating that slightly less than one-third of nestlings survive the interval between fledging and their first potential breeding season. Resighting probability was high for both sexes, although higher for adult males (0.94) than for adult female (0.84). The latter finding is consistent with the higher site fidelity of males than of females. Resighting probabilities for juveniles were much lower at 0.68 and 0.40 for males and females, respectively. Again, this was expected because natal site fidelity is typically much lower than breeding site fidelity. Compared to most other reports, resighting probability and return rates of juvenile kingbirds was high, presumably because the riparian habitats where kingbirds breed at Malheur function as an ecological island surrounded by, for kingbirds, unsuitable high desert habitat. Thus, unless they disperse very long distances, the only option for juvenile kingbirds is to begin nesting on the refuge. The collection of blood samples from birds has become an increasingly common practice in ornithology. The data that can be gained from these samples allow a number of interesting questions to be asked such as understanding the genetic mating system of a species, patterns of hormone secretion, and discerning migratory pathways via stable isotope analysis. The volume of blood collected is usually small and was assumed to cause no long-term negative effects on sampled individuals. However, few studies have rigorously examined the effect of blood sampling on survival. I used a multistate mark-recapture analysis to assess the effect blood sampling on annual survival of kingbirds by combining the annual survival data described above with whether or not individuals were subjected to blood sample collection. The results of this analysis indicated that blood sampling had no effect on annual survival rates of kingbirds. Whether or not this is the case for other species remains to be seen. However, my results support the assumption that when done correctly, blood sampling has little to no long-term negative effects on birds. Comparative analyses of many bird species show that testosterone secretion exhibits fairly predictable patterns among breeding birds of different mating systems. Monogamous species reach a peak during mate attraction and the period of female fertility which is then followed by a sharp decline when young are in the nest. By contrast, males of polygynous species tend to maintain higher levels of testosterone throughout the breeding season to, presumably, maximize opportunities for acquiring additional mates. Kingbirds are socially monogamous but cryptically polygamous. However, because of high rates of extra-pair paternity, variation in reproductive success among males is much higher than what is expected for a monogamous species, and, instead, is similar to what has been reported for polygynous species. Therefore from 2005 to 2009 I measured testosterone concentrations from the blood samples collected from male kingbirds to determine the breeding season profile of testosterone secretion in kingbirds and to understand the factors that influence testosterone variation among individuals. Contrary to expected for a monogamous species, the testosterone profile of kingbirds did not exhibit the brief peak in testosterone followed by a precipitous decline. Testosterone peaked early in the season, but declined very slowly as the nest cycle progressed. I attributed this gradual decline in testosterone to the cryptically polygynous nature of the kingbird extra-pair mating system. I also found substantial variation in testosterone concentration among male kingbirds and was able to identify several factors contributing to this variation. As expected, testosterone declined as the breeding season progressed (independent of stage in nest cycle) and nest density increased, while increases in testosterone were correlated with the number of fertile females within the population. This suggests that male kingbirds were capable of modulating testosterone concentrations to both cope with an increase in conspecific density, but also to ready themselves for times when extra-pair copulations were more likely. Close

• 2. [Article] Consequences of Mycorrhizal Colonization for Piriqueta Morphotypes under Drought Stress

  Field and greenhouse studies have shown that arbuscular mycorrhizal fungi (AMF) can improve plant growth in environments with restricted water availability. The benefits of AMF symbiosis vary among plant ...

  Citation × Citation Citation Abstract Copy Title: Consequences of Mycorrhizal Colonization for Piriqueta Morphotypes under Drought Stress Author: Sochacki, Paul, Rhode Ward, Jennifer, Cruzan, Mitchell B. Year: 2013 Field and greenhouse studies have shown that arbuscular mycorrhizal fungi (AMF) can improve plant growth in environments with restricted water availability. The benefits of AMF symbiosis vary among plant species, but the extent to which AMF-mediated drought tolerance varies among subspecific taxa remains poorly understood. In this study, we examine differences in AMF response among three recently diverged, ecologically heterogeneous plant taxa (morphotypes) within the Piriqueta cistoides spp. caroliniana complex. We performed a greenhouse experiment using cuttings of each morphotype inoculated in field-collected soil to test for inoculum source effects of AMF on plant growth under drought. Correlation between AMF colonization and plant performance under drought was significant for all three morphotypes but was strongest for viridis morphotype; this group is associated with mesic, low-phosphorous soils of south Florida slash pine flatwoods. Compared to inocula obtained from other morphotypes’ regions, the AMF obtained from one of the most arid habitats (caroliniana) promoted an equal or greater amount of growth in host plants despite relatively low levels of root colonization. These findings suggest that both genetic divergence among morphotypes and the source of AMF inoculum affect plant growth under drought in P. c. ssp. caroliniana complex. Title: Consequences of Mycorrhizal Colonization for Piriqueta Morphotypes under Drought Stress Author: Sochacki, Paul, Rhode Ward, Jennifer, Cruzan, Mitchell B. Year: 2013 Field and greenhouse studies have shown that arbuscular mycorrhizal fungi (AMF) can improve plant growth in environments with restricted water availability. The benefits of AMF symbiosis vary among plant species, but the extent to which AMF-mediated drought tolerance varies among subspecific taxa remains poorly understood. In this study, we examine differences in AMF response among three recently diverged, ecologically heterogeneous plant taxa (morphotypes) within the Piriqueta cistoides spp. caroliniana complex. We performed a greenhouse experiment using cuttings of each morphotype inoculated in field-collected soil to test for inoculum source effects of AMF on plant growth under drought. Correlation between AMF colonization and plant performance under drought was significant for all three morphotypes but was strongest for viridis morphotype; this group is associated with mesic, low-phosphorous soils of south Florida slash pine flatwoods. Compared to inocula obtained from other morphotypes’ regions, the AMF obtained from one of the most arid habitats (caroliniana) promoted an equal or greater amount of growth in host plants despite relatively low levels of root colonization. These findings suggest that both genetic divergence among morphotypes and the source of AMF inoculum affect plant growth under drought in P. c. ssp. caroliniana complex. Close

• 3. [Article] The Impeccable Timing of the Apple Maggot Fly, <i>Rhagoletis pomonella</i> (Dipetera: Tephritidae), and its Implications for Ecological Speciation

  Speciation is the process by which life diversifies into discrete forms, and understanding its underlying mechanisms remains a primary focus for biologists. Increasingly, empirical studies are helping ...

  Citation × Citation Citation Abstract Copy Title: The Impeccable Timing of the Apple Maggot Fly, <i>Rhagoletis pomonella</i> (Dipetera: Tephritidae), and its Implications for Ecological Speciation Author: Mattsson, Monte Arthur Year: 2015 Speciation is the process by which life diversifies into discrete forms, and understanding its underlying mechanisms remains a primary focus for biologists. Increasingly, empirical studies are helping explain the role of ecology in generating biodiversity. Adaptive radiations are often propelled by selective fitness tradeoffs experienced by individuals that invade new habitats, resulting in reproductive isolation from ancestral conspecifics and potentially cladogenesis. Host specialist insects are among the most speciose organisms known and serve as highly useful models for studying adaptive radiations. We are just beginning to understand the pace and degree with which these insects diversify. The apple maggot, Rhagoletis pomonella, is a well-studied insect whose eastern and southern populations are models for ecological speciation. Recently (40–65 ya), the fly has invaded the Pacific Northwestern United States through human-transported apples infested with larvae. There, populations of R. pomonella have rapidly colonized two novel hawthorn hosts whose fruiting times bracket apple’s (early-season native Crataegus douglasii and introduced C. monogyna, which fruits late in the season). The recent introduction might initiate host shifts, providing opportunities to examine the pace and mechanistic means with which host races (an evolutionary stage preceding speciation) become established. Here, I demonstrate that host-associated populations at a site in southwest Washington are partially allochronically isolated from one another, and life cycles temporally match with natal host fruit ripening times in sympatry. If spatially widespread, these temporal barriers could result in reproductive isolation and possibly cladogenesis. Implications of these findings reach beyond academic import, as R. pomonella is expanding not only its host range, but its geographic range is encroaching upon central Washington, the site of a multi-billion dollar per year apple-growing industry. Title: The Impeccable Timing of the Apple Maggot Fly, <i>Rhagoletis pomonella</i> (Dipetera: Tephritidae), and its Implications for Ecological Speciation Author: Mattsson, Monte Arthur Year: 2015 Speciation is the process by which life diversifies into discrete forms, and understanding its underlying mechanisms remains a primary focus for biologists. Increasingly, empirical studies are helping explain the role of ecology in generating biodiversity. Adaptive radiations are often propelled by selective fitness tradeoffs experienced by individuals that invade new habitats, resulting in reproductive isolation from ancestral conspecifics and potentially cladogenesis. Host specialist insects are among the most speciose organisms known and serve as highly useful models for studying adaptive radiations. We are just beginning to understand the pace and degree with which these insects diversify. The apple maggot, Rhagoletis pomonella, is a well-studied insect whose eastern and southern populations are models for ecological speciation. Recently (40–65 ya), the fly has invaded the Pacific Northwestern United States through human-transported apples infested with larvae. There, populations of R. pomonella have rapidly colonized two novel hawthorn hosts whose fruiting times bracket apple’s (early-season native Crataegus douglasii and introduced C. monogyna, which fruits late in the season). The recent introduction might initiate host shifts, providing opportunities to examine the pace and mechanistic means with which host races (an evolutionary stage preceding speciation) become established. Here, I demonstrate that host-associated populations at a site in southwest Washington are partially allochronically isolated from one another, and life cycles temporally match with natal host fruit ripening times in sympatry. If spatially widespread, these temporal barriers could result in reproductive isolation and possibly cladogenesis. Implications of these findings reach beyond academic import, as R. pomonella is expanding not only its host range, but its geographic range is encroaching upon central Washington, the site of a multi-billion dollar per year apple-growing industry. Close

• 4. [Article] Survivorship and Breeding Dispersal Patterns of a Migratory, Socially Monogamous Passerine; the Scissor-tailed Flycatcher (Tyrannus forficatus)

  Survivorship (the likelihood of survival from one year to the next) and breeding dispersal (movement between breeding seasons) exhibit considerable variability at both the inter- and intraspecific levels. ...

  Citation × Citation Citation Abstract Copy Title: Survivorship and Breeding Dispersal Patterns of a Migratory, Socially Monogamous Passerine; the Scissor-tailed Flycatcher (Tyrannus forficatus) Author: Becker, Adam John Year: 2016 Survivorship (the likelihood of survival from one year to the next) and breeding dispersal (movement between breeding seasons) exhibit considerable variability at both the inter- and intraspecific levels. Using eight years of data (2008-2015), from my study site in southwest Oklahoma, I characterized survivorship and breeding dispersal of the Scissor-tailed Flycatcher (Tyrannus forficatus) in a mixed-grass prairie ecoregion. My results suggest that estimated survivorship of Scissor-tailed Flycatchers is low, especially to the congeneric Eastern Kingbird (T. tyrannus), and was likely underestimated due to the tendency of Scissor-tailed Flycatchers to disperse long distances. Title: Survivorship and Breeding Dispersal Patterns of a Migratory, Socially Monogamous Passerine; the Scissor-tailed Flycatcher (Tyrannus forficatus) Author: Becker, Adam John Year: 2016 Survivorship (the likelihood of survival from one year to the next) and breeding dispersal (movement between breeding seasons) exhibit considerable variability at both the inter- and intraspecific levels. Using eight years of data (2008-2015), from my study site in southwest Oklahoma, I characterized survivorship and breeding dispersal of the Scissor-tailed Flycatcher (Tyrannus forficatus) in a mixed-grass prairie ecoregion. My results suggest that estimated survivorship of Scissor-tailed Flycatchers is low, especially to the congeneric Eastern Kingbird (T. tyrannus), and was likely underestimated due to the tendency of Scissor-tailed Flycatchers to disperse long distances. Close

• 5. [Article] Mechanisms of Adaptation in the Newly Invasive Species <i>Brachypodium sylvaticum</i> (Hudson) Beauv.

  It is common knowledge that invasive species cause worldwide ecological and economic damage, and are nearly impossible to eradicate. However, upon introduction to a novel environment, alien species should ...

  Citation × Citation Citation Abstract Copy Title: Mechanisms of Adaptation in the Newly Invasive Species <i>Brachypodium sylvaticum</i> (Hudson) Beauv. Author: Marchini, Gina Lola Year: 2015 It is common knowledge that invasive species cause worldwide ecological and economic damage, and are nearly impossible to eradicate. However, upon introduction to a novel environment, alien species should be the underdogs: They are present in small numbers, possess low genetic diversity, and have not adapted to the climate and competitors present in the new habitat. So, how are alien species able to invade an environment occupied by native species that have already adapted to the local environment? To discover some answers to this apparent paradox I conducted four ecological genetic studies that utilized the invasive species Brachypodium sylvaticum (Hudson) Beauv. to determine mechanisms contributing to adaptation and success in the novel habitat. The first study used simulations and experiments to test the hypothesis that genetic purging, the process where genetic load is reduced by selection against the recessive deleterious alleles expressed in the homozygous state, promotes invasive range expansion. I found that homozygous populations on B. sylvaticum's range periphery displayed lower inbreeding depression compared to heterozygous populations near introduction sites. Empirical tests with B. sylvaticum further demonstrate that purging of genetic load is a plausible scenario promoting range expansion during invasion. Next, I explored how the interaction between population genetic diversity and the environment contributed to the establishment and spread of Brachypodium sylvaticum. I found that nitrogen application increases both final size and shoot biomass for B. sylvaticum individuals from source populations with low HS levels to levels found in individuals from populations with high HS. A coefficient of relative competition intensity index (RCI) displayed reduced effects of interspecific competition on B. sylvaticum biomass in high nitrogen plots. Results show that elevated nitrogen deposition is a factor that increases establishment of introduced species with historically small effective population sizes. Thirdly, I investigated phenotypic differentiation during the establishment and range expansion of Brachypodium sylvaticum. Utilizing a novel approach, unique alleles were used to determine the genetic probability of contribution from native source regions to invasive regions. These probabilities were integrated into QST-FST comparisons to determine the influence of selection and genetic drift on twelve physiological and anatomical traits associated with drought stress. Phenotypic divergence greater than neutral expectations was found for five traits between native and invasive populations, indicating selective divergence. Results from this study show that the majority of divergence in B. sylvaticum occurred after introduction to the novel environment, but prior to invasive range expansion. The final chapter of my dissertation investigates the adaptive role of genetic differentiation and plasticity for Brachypodium sylvaticum invasion. Plasticity was measured across treatments of contrasting water availability. Linear and nonlinear selection gradients determined the effect of directional and quadratic selection on plasticity and genetic differentiation. Invasive trait divergence was a consequence of post-introduction selection leading to genetic differentiation, as there were no plastic responses to contrasting water availability for any measured traits. Genetic divergence of invasive plants was not consistently in the direction indicated by selection, suggesting limitations of selection that may be a consequence of physical constraints and/or tradeoffs between growth and abiotic tolerance. Results suggest that selection, rather than plasticity, is driving phenotypic change in the invaded environment. The combined volume of these studies contributes significantly to the field of invasion and plant biology by providing novel insights into the processes underlying range expansion, adaptation, and ultimately, evolution of introduced species. Title: Mechanisms of Adaptation in the Newly Invasive Species <i>Brachypodium sylvaticum</i> (Hudson) Beauv. Author: Marchini, Gina Lola Year: 2015 It is common knowledge that invasive species cause worldwide ecological and economic damage, and are nearly impossible to eradicate. However, upon introduction to a novel environment, alien species should be the underdogs: They are present in small numbers, possess low genetic diversity, and have not adapted to the climate and competitors present in the new habitat. So, how are alien species able to invade an environment occupied by native species that have already adapted to the local environment? To discover some answers to this apparent paradox I conducted four ecological genetic studies that utilized the invasive species Brachypodium sylvaticum (Hudson) Beauv. to determine mechanisms contributing to adaptation and success in the novel habitat. The first study used simulations and experiments to test the hypothesis that genetic purging, the process where genetic load is reduced by selection against the recessive deleterious alleles expressed in the homozygous state, promotes invasive range expansion. I found that homozygous populations on B. sylvaticum's range periphery displayed lower inbreeding depression compared to heterozygous populations near introduction sites. Empirical tests with B. sylvaticum further demonstrate that purging of genetic load is a plausible scenario promoting range expansion during invasion. Next, I explored how the interaction between population genetic diversity and the environment contributed to the establishment and spread of Brachypodium sylvaticum. I found that nitrogen application increases both final size and shoot biomass for B. sylvaticum individuals from source populations with low HS levels to levels found in individuals from populations with high HS. A coefficient of relative competition intensity index (RCI) displayed reduced effects of interspecific competition on B. sylvaticum biomass in high nitrogen plots. Results show that elevated nitrogen deposition is a factor that increases establishment of introduced species with historically small effective population sizes. Thirdly, I investigated phenotypic differentiation during the establishment and range expansion of Brachypodium sylvaticum. Utilizing a novel approach, unique alleles were used to determine the genetic probability of contribution from native source regions to invasive regions. These probabilities were integrated into QST-FST comparisons to determine the influence of selection and genetic drift on twelve physiological and anatomical traits associated with drought stress. Phenotypic divergence greater than neutral expectations was found for five traits between native and invasive populations, indicating selective divergence. Results from this study show that the majority of divergence in B. sylvaticum occurred after introduction to the novel environment, but prior to invasive range expansion. The final chapter of my dissertation investigates the adaptive role of genetic differentiation and plasticity for Brachypodium sylvaticum invasion. Plasticity was measured across treatments of contrasting water availability. Linear and nonlinear selection gradients determined the effect of directional and quadratic selection on plasticity and genetic differentiation. Invasive trait divergence was a consequence of post-introduction selection leading to genetic differentiation, as there were no plastic responses to contrasting water availability for any measured traits. Genetic divergence of invasive plants was not consistently in the direction indicated by selection, suggesting limitations of selection that may be a consequence of physical constraints and/or tradeoffs between growth and abiotic tolerance. Results suggest that selection, rather than plasticity, is driving phenotypic change in the invaded environment. The combined volume of these studies contributes significantly to the field of invasion and plant biology by providing novel insights into the processes underlying range expansion, adaptation, and ultimately, evolution of introduced species. Close

• 6. [Article] Green Roofs and Urban Biodiversity: Their Role as Invertebrate Habitat and the Effect of Design on Beetle Community

  With over half the world's population now living in cities, urban areas represent one of earth's few ecosystems that are increasing in extent, and are sites of altered biogeochemical cycles, habitat fragmentation, ...

  Citation × Citation Citation Abstract Copy Title: Green Roofs and Urban Biodiversity: Their Role as Invertebrate Habitat and the Effect of Design on Beetle Community Author: Gonsalves, Sydney Marie Year: 2016 With over half the world's population now living in cities, urban areas represent one of earth's few ecosystems that are increasing in extent, and are sites of altered biogeochemical cycles, habitat fragmentation, and changes in biodiversity. However, urban green spaces, including green roofs, can also provide important pools of biodiversity and contribute to regional gamma diversity, while novel species assemblages can enhance some ecosystem services. Green roofs may also mitigate species loss in urban areas and have been shown to support a surprising diversity of invertebrates, including rare and endangered species. In the first part of this study I reviewed the literature on urban invertebrate communities and diversity to better understand the role of green roofs in providing habitat in the context of the larger urban mosaic. My review concluded that, while other factors such as surrounding land use and connectivity are also important to specific invertebrate taxa, local habitat variables contribute substantially to the structure and diversity of urban invertebrate communities. The importance of local habitat variables in urban green spaces and strong support for the habitat complexity hypothesis in a number of other ecosystems has led to proposals that "biodiverse" roofs--those intentionally designed with varied substrate depth, greater plant diversity, or added elements such as logs or stones--would support greater invertebrate diversity, but there is currently limited peer reviewed data to support this. In order to address the habitat complexity hypothesis in the context of green roofs, in the second part of this study I surveyed three roofs designed primarily for stormwater management, three biodiverse roofs, and five ground-level green spaces, from March until September of 2014 in the Portland metropolitan area. Beetles (Coleoptera) were sampled bi-weekly as representatives of total species diversity. Biodiverse roofs had greater richness, abundance, and diversity of beetle species compared to stormwater roofs, but were not more diverse than ground sites. Both biodiverse roofs and ground sites had approximately 20% native beetle species while stormwater roofs had only 5%. Functional diversity was also higher on biodiverse roofs with an average of seven trophic groups represented, while stormwater roofs averaged only three. Ground sites, biodiverse roofs, and stormwater roofs each grouped distinctively in terms of beetle community composition and biodiverse roof communities were found to be positively correlated with roof age, percent plant cover, average plant height, and plant species richness. These results support the findings of previous studies on the importance of local variables in structuring urban invertebrate communities and suggest that biodiverse design can reliably increase greenroof diversity, with the caution that they remain no replacement for ground level conservation. Title: Green Roofs and Urban Biodiversity: Their Role as Invertebrate Habitat and the Effect of Design on Beetle Community Author: Gonsalves, Sydney Marie Year: 2016 With over half the world's population now living in cities, urban areas represent one of earth's few ecosystems that are increasing in extent, and are sites of altered biogeochemical cycles, habitat fragmentation, and changes in biodiversity. However, urban green spaces, including green roofs, can also provide important pools of biodiversity and contribute to regional gamma diversity, while novel species assemblages can enhance some ecosystem services. Green roofs may also mitigate species loss in urban areas and have been shown to support a surprising diversity of invertebrates, including rare and endangered species. In the first part of this study I reviewed the literature on urban invertebrate communities and diversity to better understand the role of green roofs in providing habitat in the context of the larger urban mosaic. My review concluded that, while other factors such as surrounding land use and connectivity are also important to specific invertebrate taxa, local habitat variables contribute substantially to the structure and diversity of urban invertebrate communities. The importance of local habitat variables in urban green spaces and strong support for the habitat complexity hypothesis in a number of other ecosystems has led to proposals that "biodiverse" roofs--those intentionally designed with varied substrate depth, greater plant diversity, or added elements such as logs or stones--would support greater invertebrate diversity, but there is currently limited peer reviewed data to support this. In order to address the habitat complexity hypothesis in the context of green roofs, in the second part of this study I surveyed three roofs designed primarily for stormwater management, three biodiverse roofs, and five ground-level green spaces, from March until September of 2014 in the Portland metropolitan area. Beetles (Coleoptera) were sampled bi-weekly as representatives of total species diversity. Biodiverse roofs had greater richness, abundance, and diversity of beetle species compared to stormwater roofs, but were not more diverse than ground sites. Both biodiverse roofs and ground sites had approximately 20% native beetle species while stormwater roofs had only 5%. Functional diversity was also higher on biodiverse roofs with an average of seven trophic groups represented, while stormwater roofs averaged only three. Ground sites, biodiverse roofs, and stormwater roofs each grouped distinctively in terms of beetle community composition and biodiverse roof communities were found to be positively correlated with roof age, percent plant cover, average plant height, and plant species richness. These results support the findings of previous studies on the importance of local variables in structuring urban invertebrate communities and suggest that biodiverse design can reliably increase greenroof diversity, with the caution that they remain no replacement for ground level conservation. Close

• 7. [Article] Nuwuvi (Southern Paiute) Ecological Knowledge of Piñon-Juniper Woodlands: Implications for Conservation and Sustainable Resource Use in Two Southern Nevada Protected Areas

  Nuwuvi (Southern Paiute) have inhabited the southern Great Basin for thousands of years, and consider Nuvagantu (where snow sits) in the Spring Mountains landscape to be the locus of their creation as ...

  Citation × Citation Citation Abstract Copy Title: Nuwuvi (Southern Paiute) Ecological Knowledge of Piñon-Juniper Woodlands: Implications for Conservation and Sustainable Resource Use in Two Southern Nevada Protected Areas Author: Lefler, Brian John Year: 2014 Nuwuvi (Southern Paiute) have inhabited the southern Great Basin for thousands of years, and consider Nuvagantu (where snow sits) in the Spring Mountains landscape to be the locus of their creation as a people. Their ancestral territory spans parts of Nevada, Utah, Arizona, and California. My research identifies and describes the heterogeneous character of Nuwuvi ecological knowledge (NEK) of piñon-juniper woodland ecosystems within two federal protected areas (PAs) in southeastern Nevada, the Spring Mountains National Recreation Area (SMNRA) and the Desert National Wildlife Refuge (DNWR), as remembered and practiced to varying degrees by 22 select Nuwuvi knowledge holders. I focus my investigation on four primary aspects of NEK. First, drawing from data obtained through ethnoecological research, I discuss how Nuwuvi ecological knowledge evolved through protracted observation and learning from past resource depletions, and adapted to various environmental and socio-economic drivers of change induced since Euro-American incursion. Second, I argue that Nuwuvi management practices operate largely within a framework of non-equilibrium ecology, marked by low to intermediate disturbances and guided by Nuwuvi conceptions of environmental health and balance. These practices favor landscape heterogeneity and patchiness, and engender ecosystem renewal, expanded ecotones, and increased biodiversity. I then consider the third and fourth aspects of NEK as two case studies that consider NEK at the individual, species, population, habitat, and landscape scales. These case studies operationalize NEK as a relevant body of knowledge and techniques conducive to collaborative resource stewardship initiatives with federal land management agency partners. In the first case study I suggest that the Great Basin piñon pines are Nuwuvi cultural keystone species (CKS), evaluating their central importance to Nuwuvi according to several criteria including number of uses, role in ritual and story, and uniqueness relative to other species. In the second case study I contend that local social institutions regulated Nuwuvi resource use in the past and in some cases continued to do so at the time of study. These local social institutions included a system of resource extraction and habitat entrance taboos that may have mitigated impacts and supported sustainable resource use and conservation. The implications of this research are that Nuwuvi ecological knowledge, disturbance-based adaptive management practices, and resource and habitat taboos are relevant to contemporary land management concerns in piñon-juniper woodlands, offering complementary approaches to adaptive management as practiced in the SMNRA and the DNWR despite divergent epistemological foundations. My research contributed to the Nuwuvi Knowledge-to-Action Project, an applied government-to-government consultation, collaborative resource stewardship, and cultural revitalization project facilitated by The Mountain Institute among seven Nuwuvi Nations, the U.S. Forest Service, and the U.S. Fish and Wildlife Service. Title: Nuwuvi (Southern Paiute) Ecological Knowledge of Piñon-Juniper Woodlands: Implications for Conservation and Sustainable Resource Use in Two Southern Nevada Protected Areas Author: Lefler, Brian John Year: 2014 Nuwuvi (Southern Paiute) have inhabited the southern Great Basin for thousands of years, and consider Nuvagantu (where snow sits) in the Spring Mountains landscape to be the locus of their creation as a people. Their ancestral territory spans parts of Nevada, Utah, Arizona, and California. My research identifies and describes the heterogeneous character of Nuwuvi ecological knowledge (NEK) of piñon-juniper woodland ecosystems within two federal protected areas (PAs) in southeastern Nevada, the Spring Mountains National Recreation Area (SMNRA) and the Desert National Wildlife Refuge (DNWR), as remembered and practiced to varying degrees by 22 select Nuwuvi knowledge holders. I focus my investigation on four primary aspects of NEK. First, drawing from data obtained through ethnoecological research, I discuss how Nuwuvi ecological knowledge evolved through protracted observation and learning from past resource depletions, and adapted to various environmental and socio-economic drivers of change induced since Euro-American incursion. Second, I argue that Nuwuvi management practices operate largely within a framework of non-equilibrium ecology, marked by low to intermediate disturbances and guided by Nuwuvi conceptions of environmental health and balance. These practices favor landscape heterogeneity and patchiness, and engender ecosystem renewal, expanded ecotones, and increased biodiversity. I then consider the third and fourth aspects of NEK as two case studies that consider NEK at the individual, species, population, habitat, and landscape scales. These case studies operationalize NEK as a relevant body of knowledge and techniques conducive to collaborative resource stewardship initiatives with federal land management agency partners. In the first case study I suggest that the Great Basin piñon pines are Nuwuvi cultural keystone species (CKS), evaluating their central importance to Nuwuvi according to several criteria including number of uses, role in ritual and story, and uniqueness relative to other species. In the second case study I contend that local social institutions regulated Nuwuvi resource use in the past and in some cases continued to do so at the time of study. These local social institutions included a system of resource extraction and habitat entrance taboos that may have mitigated impacts and supported sustainable resource use and conservation. The implications of this research are that Nuwuvi ecological knowledge, disturbance-based adaptive management practices, and resource and habitat taboos are relevant to contemporary land management concerns in piñon-juniper woodlands, offering complementary approaches to adaptive management as practiced in the SMNRA and the DNWR despite divergent epistemological foundations. My research contributed to the Nuwuvi Knowledge-to-Action Project, an applied government-to-government consultation, collaborative resource stewardship, and cultural revitalization project facilitated by The Mountain Institute among seven Nuwuvi Nations, the U.S. Forest Service, and the U.S. Fish and Wildlife Service. Close

• 8. [Article] Small Tidal Channels Improve Foraging Opportunities for Calidris Shorebirds

  Estuarine intertidal habitats are heterogeneous, therefore migratory shorebirds are expected to forage in microhabitats where they can maximize their energy intake. Identifying proximate factors that migratory ...

  Citation × Citation Citation Abstract Copy Title: Small Tidal Channels Improve Foraging Opportunities for Calidris Shorebirds Author: Miller, Aileen K., de Rivera, Catherine E. Year: 2014 Estuarine intertidal habitats are heterogeneous, therefore migratory shorebirds are expected to forage in microhabitats where they can maximize their energy intake. Identifying proximate factors that migratory shorebirds use to accept or reject a particular habitat patch will help land managers make conservation and restoration decisions that provide the greatest benefits to shorebird populations during migration, a period of intense energy usage. We examined whether small semipermanent tidal channels were preferentially used by foraging Western Sandpipers (Calidris mauri) and Dunlins (C. alpina) during a spring migratory stopover in Bandon Marsh, an Oregon, USA, estuary. Further, we tested alternative hypotheses about how channels might be beneficial to shorebirds by examining infauna abundance, composition, and depth alongside channels compared with that in open mudflats. The densities of Western Sandpipers and Dunlins were at least 4 times higher along channels than in open mudflat habitats. Infauna along channels were twice as abundant as those in the surrounding mudflats, and infauna were generally found closer to the surface. Furthermore, sediment alongside channels required less than half the force to probe, probably increasing shorebirds' overall access to prey. We conclude that shorebirds selected microhabitats near channels because of the greater food abundance and availability. Additional evidence suggested that another microhabitat, patches of red algae (Gracilaria sp.), may also have conferred advantages on foraging shorebirds. Channels may be a valuable microhabitat to consider when selecting conservation areas for shorebirds, and maintaining channels (e.g., through maintaining natural hydrology and sedimentation patterns) should be considered in management practices. Title: Small Tidal Channels Improve Foraging Opportunities for Calidris Shorebirds Author: Miller, Aileen K., de Rivera, Catherine E. Year: 2014 Estuarine intertidal habitats are heterogeneous, therefore migratory shorebirds are expected to forage in microhabitats where they can maximize their energy intake. Identifying proximate factors that migratory shorebirds use to accept or reject a particular habitat patch will help land managers make conservation and restoration decisions that provide the greatest benefits to shorebird populations during migration, a period of intense energy usage. We examined whether small semipermanent tidal channels were preferentially used by foraging Western Sandpipers (Calidris mauri) and Dunlins (C. alpina) during a spring migratory stopover in Bandon Marsh, an Oregon, USA, estuary. Further, we tested alternative hypotheses about how channels might be beneficial to shorebirds by examining infauna abundance, composition, and depth alongside channels compared with that in open mudflats. The densities of Western Sandpipers and Dunlins were at least 4 times higher along channels than in open mudflat habitats. Infauna along channels were twice as abundant as those in the surrounding mudflats, and infauna were generally found closer to the surface. Furthermore, sediment alongside channels required less than half the force to probe, probably increasing shorebirds' overall access to prey. We conclude that shorebirds selected microhabitats near channels because of the greater food abundance and availability. Additional evidence suggested that another microhabitat, patches of red algae (Gracilaria sp.), may also have conferred advantages on foraging shorebirds. Channels may be a valuable microhabitat to consider when selecting conservation areas for shorebirds, and maintaining channels (e.g., through maintaining natural hydrology and sedimentation patterns) should be considered in management practices. Close

• 9. [Article] Accelerated Erosion of Saltmarshes Infested by the Non-Native Burrowing Crustacean Sphaeroma Quoianum

  Lateral erosion of saltmarshes is affected by many abiotic and biotic factors. While abiotic factors are typically regarded as primary drivers of erosion, biotic influences such as burrowing or bioturbating ...

  Citation × Citation Citation Abstract Copy Title: Accelerated Erosion of Saltmarshes Infested by the Non-Native Burrowing Crustacean Sphaeroma Quoianum Author: Davidson, Timothy Mathias, de Rivera, Catherine E. Year: 2010 Lateral erosion of saltmarshes is affected by many abiotic and biotic factors. While abiotic factors are typically regarded as primary drivers of erosion, biotic influences such as burrowing or bioturbating taxa can also extensively modify the physical structure of this marine habitat. Many estuaries on the Pacific coast of North America have been invaded by populations of the burrowing non-native isopod Sphaeroma quoianum, which are thought to exacerbate the erosion of saltmarshes. We conducted a mensurative experiment to examine the relationship between populations of S. quoianum and lateral erosion rate of saltmarshes in Coos Bay, Oregon, USA. After 1 yr, we measured higher lateral erosion rates, more undercutting, and higher numbers of calved and slumped marsh sections in marsh sites infested by S. quoianum than in uninfested sites. This effect was also consistent at smaller spatial scales: we found lateral erosion was 300% higher in burrowed areas than in adjacent (within approx. 1 m) unburrowed areas within infested sites. Our results suggest that the prodigious burrowing by dense populations of S. quoianum can accelerate erosion rates in saltmarshes and are substantial geomorphic agents of change in estuaries on the Pacific coast of North America. Title: Accelerated Erosion of Saltmarshes Infested by the Non-Native Burrowing Crustacean Sphaeroma Quoianum Author: Davidson, Timothy Mathias, de Rivera, Catherine E. Year: 2010 Lateral erosion of saltmarshes is affected by many abiotic and biotic factors. While abiotic factors are typically regarded as primary drivers of erosion, biotic influences such as burrowing or bioturbating taxa can also extensively modify the physical structure of this marine habitat. Many estuaries on the Pacific coast of North America have been invaded by populations of the burrowing non-native isopod Sphaeroma quoianum, which are thought to exacerbate the erosion of saltmarshes. We conducted a mensurative experiment to examine the relationship between populations of S. quoianum and lateral erosion rate of saltmarshes in Coos Bay, Oregon, USA. After 1 yr, we measured higher lateral erosion rates, more undercutting, and higher numbers of calved and slumped marsh sections in marsh sites infested by S. quoianum than in uninfested sites. This effect was also consistent at smaller spatial scales: we found lateral erosion was 300% higher in burrowed areas than in adjacent (within approx. 1 m) unburrowed areas within infested sites. Our results suggest that the prodigious burrowing by dense populations of S. quoianum can accelerate erosion rates in saltmarshes and are substantial geomorphic agents of change in estuaries on the Pacific coast of North America. Close

• 10. [Article] Northern Range Expansion and Coastal Occurrences of the New Zealand Mud Snail Potamopyrgus Antipodarum (Gray, 1843) in the Northeast Pacific

  The New Zealand mud snail Potamopyrgus antipodarum (Gray, 1843) is a common invasive species in fresh and brackish water ecosystems in Europe, Australia, Japan, and North America. In some invaded habitats, ...

  Citation × Citation Citation Abstract Copy Title: Northern Range Expansion and Coastal Occurrences of the New Zealand Mud Snail Potamopyrgus Antipodarum (Gray, 1843) in the Northeast Pacific Author: Davidson, Timothy Mathias, Brenneis, Valance E. F., de Rivera, Catherine E., Draheim, Robyn, Gillespie, Graham E. Year: 2008 The New Zealand mud snail Potamopyrgus antipodarum (Gray, 1843) is a common invasive species in fresh and brackish water ecosystems in Europe, Australia, Japan, and North America. In some invaded habitats, P. antipodarum can reach high densities (over 500,000 snails m-2) and dominate the biomass of the benthos, leading to detrimental impacts to native biota and changes in ecosystem dynamics. We report the previously unpublished occurrence of P. antipodarum in thirteen fresh and brackish water systems adjacent to the Pacific coast of North America including a new northern range for P. antipodarum: Port Alberni, Vancouver Island, British Columbia, Canada (49.2479º, -124.8395º). We hypothesize the snail was spread from the Columbia River Estuary to Port Alberni via recreational watercraft or infected fishing equipment. Its discovery in Port Alberni reveals the potential for other aquatic nuisance species in the lower Columbia River to spread to British Columbia. Resource managers on the Pacific coast should remain vigilant and educate the public to prevent the further spread of the P. antipodarum as well as other aquatic invaders. Title: Northern Range Expansion and Coastal Occurrences of the New Zealand Mud Snail Potamopyrgus Antipodarum (Gray, 1843) in the Northeast Pacific Author: Davidson, Timothy Mathias, Brenneis, Valance E. F., de Rivera, Catherine E., Draheim, Robyn, Gillespie, Graham E. Year: 2008 The New Zealand mud snail Potamopyrgus antipodarum (Gray, 1843) is a common invasive species in fresh and brackish water ecosystems in Europe, Australia, Japan, and North America. In some invaded habitats, P. antipodarum can reach high densities (over 500,000 snails m-2) and dominate the biomass of the benthos, leading to detrimental impacts to native biota and changes in ecosystem dynamics. We report the previously unpublished occurrence of P. antipodarum in thirteen fresh and brackish water systems adjacent to the Pacific coast of North America including a new northern range for P. antipodarum: Port Alberni, Vancouver Island, British Columbia, Canada (49.2479º, -124.8395º). We hypothesize the snail was spread from the Columbia River Estuary to Port Alberni via recreational watercraft or infected fishing equipment. Its discovery in Port Alberni reveals the potential for other aquatic nuisance species in the lower Columbia River to spread to British Columbia. Resource managers on the Pacific coast should remain vigilant and educate the public to prevent the further spread of the P. antipodarum as well as other aquatic invaders. Close