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• 11. [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

• 12. [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

• 13. [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

• 14. [Article] Biotic Resistance to Invasion: Native Predator Limits Abundance and Distribution of an Introduced Crab

  Introduced species frequently escape the natural enemies (predators, competitors, and parasites) that limit their distribution and abundance in the native range. This reduction in native predators, competitors, ...

  Citation × Citation Citation Abstract Copy Title: Biotic Resistance to Invasion: Native Predator Limits Abundance and Distribution of an Introduced Crab Author: de Rivera, Catherine E., Ruiz, Greg M., Hines, Anson, Jivoff, Paul Year: 2005 Introduced species frequently escape the natural enemies (predators, competitors, and parasites) that limit their distribution and abundance in the native range. This reduction in native predators, competitors, and parasites may result in ecological release in the introduced range. However, biological interactions also can limit the establishment and spread of nonnative populations. The extent to which such biotic resistance occurs is poorly resolved, especially for marine ecosystems. Here we test whether a native predator, the blue crab Callinectes sapidus, affects the abundance and geographic range of the introduced European green crab Carcinus maenas in eastern North America. Both crab species occur in shallow, soft-sediment habitats of bays and estuaries, and their ranges overlap in eastern North America. First, we tested for a negative relationship in the abundances of the two species from trap samples across a 640-km (5.78° latitude) coastal transect. Second, we estimated variation in predation pressure on tethered Carcinus maenas across latitude and as a function of Callinectes sapidus abundance. Third, we measured predation rates on Carcinus maenas by Callinectes sapidus in field and laboratory experiments. Our results support the hypothesis that the native predator Callinectes sapidus provides biotic resistance to invasion and prevents the southward spread and establishment of Carcinus maenas. Within and across bays, Carcinus maenas were significantly less abundant at sites and depths with Callinectes sapidus compared with areas lacking Callinectes sapidus. Moreover, no Carcinus maenas were found in Chesapeake Bay, where Callinectes sapidus were most abundant. Predation of tethered Carcinus maenas increased with Callinectes sapidus abundance. In laboratory and field experiments, Callinectes sapidus preyed readily on Carcinus maenas. Thus, we conclude the predation by Callinectes sapidus, alone or in combination with other factors, limits the abundance and geographic range of an invasive marine species. Title: Biotic Resistance to Invasion: Native Predator Limits Abundance and Distribution of an Introduced Crab Author: de Rivera, Catherine E., Ruiz, Greg M., Hines, Anson, Jivoff, Paul Year: 2005 Introduced species frequently escape the natural enemies (predators, competitors, and parasites) that limit their distribution and abundance in the native range. This reduction in native predators, competitors, and parasites may result in ecological release in the introduced range. However, biological interactions also can limit the establishment and spread of nonnative populations. The extent to which such biotic resistance occurs is poorly resolved, especially for marine ecosystems. Here we test whether a native predator, the blue crab Callinectes sapidus, affects the abundance and geographic range of the introduced European green crab Carcinus maenas in eastern North America. Both crab species occur in shallow, soft-sediment habitats of bays and estuaries, and their ranges overlap in eastern North America. First, we tested for a negative relationship in the abundances of the two species from trap samples across a 640-km (5.78° latitude) coastal transect. Second, we estimated variation in predation pressure on tethered Carcinus maenas across latitude and as a function of Callinectes sapidus abundance. Third, we measured predation rates on Carcinus maenas by Callinectes sapidus in field and laboratory experiments. Our results support the hypothesis that the native predator Callinectes sapidus provides biotic resistance to invasion and prevents the southward spread and establishment of Carcinus maenas. Within and across bays, Carcinus maenas were significantly less abundant at sites and depths with Callinectes sapidus compared with areas lacking Callinectes sapidus. Moreover, no Carcinus maenas were found in Chesapeake Bay, where Callinectes sapidus were most abundant. Predation of tethered Carcinus maenas increased with Callinectes sapidus abundance. In laboratory and field experiments, Callinectes sapidus preyed readily on Carcinus maenas. Thus, we conclude the predation by Callinectes sapidus, alone or in combination with other factors, limits the abundance and geographic range of an invasive marine species. Close

• 15. [Article] Invasive Predator, Bythotrephes, has Varied Effects on Ecosystem Function in Freshwater Lakes

  Bythotrephes longimanus is an invertebrate predator that has invaded the North American Great Lakes and a number of inland lakes, where it preys on crustacean zooplankton. We examined the effect of Bythotrephes ...

  Citation × Citation Citation Abstract Copy Title: Invasive Predator, Bythotrephes, has Varied Effects on Ecosystem Function in Freshwater Lakes Author: Strecker, Angela L., Arnott, Shelley E. Year: 2008 Bythotrephes longimanus is an invertebrate predator that has invaded the North American Great Lakes and a number of inland lakes, where it preys on crustacean zooplankton. We examined the effect of Bythotrephes on two measures of ecosystem function during a four month observational study of freshwater lakes on the boreal shield. Bythotrephes-invaded lakes had significantly lower epilimnetic zooplankton abundance and production compared to reference lakes. On average, Bythotrephes consumed 34% ofzooplankton production when it was present in lakes. There was some evidence of changes in the timing of zooplankton production, as well as shifts to cooler, less productive habitats, which may lessen the overall effect of the invader on the transfer of energy to higher trophic levels. We experimentally demonstrated a weak trophic cascade where invader predation reduced zooplankton biomass, and subsequently increased phytoplankton growth. However, the response was small in magnitude and not biologically relevant at the whole lake- scale. The most conspicuous effect of Bythotrephes that we measured was a diversion of energy away from native predators at higher trophic levels. Title: Invasive Predator, Bythotrephes, has Varied Effects on Ecosystem Function in Freshwater Lakes Author: Strecker, Angela L., Arnott, Shelley E. Year: 2008 Bythotrephes longimanus is an invertebrate predator that has invaded the North American Great Lakes and a number of inland lakes, where it preys on crustacean zooplankton. We examined the effect of Bythotrephes on two measures of ecosystem function during a four month observational study of freshwater lakes on the boreal shield. Bythotrephes-invaded lakes had significantly lower epilimnetic zooplankton abundance and production compared to reference lakes. On average, Bythotrephes consumed 34% ofzooplankton production when it was present in lakes. There was some evidence of changes in the timing of zooplankton production, as well as shifts to cooler, less productive habitats, which may lessen the overall effect of the invader on the transfer of energy to higher trophic levels. We experimentally demonstrated a weak trophic cascade where invader predation reduced zooplankton biomass, and subsequently increased phytoplankton growth. However, the response was small in magnitude and not biologically relevant at the whole lake- scale. The most conspicuous effect of Bythotrephes that we measured was a diversion of energy away from native predators at higher trophic levels. Close

• 16. [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

• 17. [Article] Complex Interactions Between Regional Dispersal of Native Taxa and an Invasive Species

  In the event of an environmental disturbance, dispersal of native taxa may provide species and genetic diversity to ecosystems, increasing the likelihood that there will be species and genotypes present ...

  Citation × Citation Citation Abstract Copy Title: Complex Interactions Between Regional Dispersal of Native Taxa and an Invasive Species Author: Strecker, Angela L., Arnott, Shelley E. Year: 2010 In the event of an environmental disturbance, dispersal of native taxa may provide species and genetic diversity to ecosystems, increasing the likelihood that there will be species and genotypes present that are less vulnerable to the disturbance. This may allow communities to maintain functioning during a disturbance and may be particularly important when the perturbation is novel to the system, such as the establishment of an invasive species. We examined how dispersal of native species may influence crustacean zooplankton communities in freshwater lakes invaded by the invertebrate predator, Bythotrephes longimanus. Using large enclosures, we experimentally tested the effect of dispersal on zooplankton community abundance, richness, and composition in (1) a community invaded by Bythotrephes, (2) the same community with the invader removed, and (3) a community that was never invaded. Dispersal increased zooplankton community abundance and richness; however, these effects were usually only significant in the invader-removed treatment. Dispersal tended to make the invader-removed communities more similar to never-invaded communities in abundance, richness, and composition. Dispersal had little effect on zooplankton abundance in the invaded community; however, richness significantly increased, and the community composition changed to resemble a never-invaded community by the end of the experiment. Our results have implications for understanding the role of dispersal during transitory states in communities. Dispersal of native taxa may be particularly important during the period between the arrival and broad-scale establishment of Bythotrephes, as dispersal through space or time (i.e., from resting eggs) may rapidly increase zooplankton abundance when the invader is absent or in low abundances. Overall, our results suggest that communities with strong local predatory and competitive interactions may be closed to immigration from colonists, but that invasive species may alter the conditions under which species can establish. These results have implications for the interaction of native and invasive species across broad spatial scales, as regional dispersal of native taxa may forestall the local extirpation of native species. In particular, transient phases that result from variable persistence of invaders within habitats or across a region may permit native colonists to successfully establish, and thus increase local and regional resistance to future disturbance. Title: Complex Interactions Between Regional Dispersal of Native Taxa and an Invasive Species Author: Strecker, Angela L., Arnott, Shelley E. Year: 2010 In the event of an environmental disturbance, dispersal of native taxa may provide species and genetic diversity to ecosystems, increasing the likelihood that there will be species and genotypes present that are less vulnerable to the disturbance. This may allow communities to maintain functioning during a disturbance and may be particularly important when the perturbation is novel to the system, such as the establishment of an invasive species. We examined how dispersal of native species may influence crustacean zooplankton communities in freshwater lakes invaded by the invertebrate predator, Bythotrephes longimanus. Using large enclosures, we experimentally tested the effect of dispersal on zooplankton community abundance, richness, and composition in (1) a community invaded by Bythotrephes, (2) the same community with the invader removed, and (3) a community that was never invaded. Dispersal increased zooplankton community abundance and richness; however, these effects were usually only significant in the invader-removed treatment. Dispersal tended to make the invader-removed communities more similar to never-invaded communities in abundance, richness, and composition. Dispersal had little effect on zooplankton abundance in the invaded community; however, richness significantly increased, and the community composition changed to resemble a never-invaded community by the end of the experiment. Our results have implications for understanding the role of dispersal during transitory states in communities. Dispersal of native taxa may be particularly important during the period between the arrival and broad-scale establishment of Bythotrephes, as dispersal through space or time (i.e., from resting eggs) may rapidly increase zooplankton abundance when the invader is absent or in low abundances. Overall, our results suggest that communities with strong local predatory and competitive interactions may be closed to immigration from colonists, but that invasive species may alter the conditions under which species can establish. These results have implications for the interaction of native and invasive species across broad spatial scales, as regional dispersal of native taxa may forestall the local extirpation of native species. In particular, transient phases that result from variable persistence of invaders within habitats or across a region may permit native colonists to successfully establish, and thus increase local and regional resistance to future disturbance. Close