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• 1. [Article] The physiology ecology and run diversity of adult Pacific lamprey, Entosphenus tridentatus, during the freshwater spawning migration

  Pacific lamprey, Entosphenus tridentatus, have shown recent and rapid declines in abundance. These anadromous fish return to streams where they mature, spawn and die. It has been inferred that Pacific ...

  Citation × Citation Citation Abstract Copy Title: The physiology ecology and run diversity of adult Pacific lamprey, Entosphenus tridentatus, during the freshwater spawning migration Author: Clemens, Benjamin Jacob, 1976- Pacific lamprey, Entosphenus tridentatus, have shown recent and rapid declines in abundance. These anadromous fish return to streams where they mature, spawn and die. It has been inferred that Pacific lamprey enter freshwater and reside for ~ 1 year before spawning. This long exposure to the freshwater environment may affect the plasticity of the maturation process and the migration timing of Pacific lamprey. Diversity in run times and body size has been observed for Pacific lamprey, yet it is unknown if this diversity is induced by the freshwater environment or if it is genetic. My first goal was to describe the maturation and migration characteristics of adult Pacific lamprey during their freshwater migration. My second goal was to use these data to make an estimation of the run diversity in Pacific lamprey. I conducted three complementary studies, in the laboratory and the field, to achieve these goals. I held immature adult lamprey (non-ripe fish that had ceased parasitic feeding in the ocean and had returned to freshwater) in the laboratory at temperatures that mimicked what these fish would experience in the wild, during the summer (mean: 21.8 °C), and another group of lamprey at cooler temperature (mean: 13.6 °C) to compare maturation timing and characteristics. The warm water group of lamprey showed significantly greater proportional decreases in body mass following temperature exposure than fish in the cooler water. All fish exposed to the warm water matured the following spring (8-10 months later) whereas only about half of the fish from the cool water exposure matured. To understand the migration distances and timing of adult Pacific lamprey, I tracked radio-tagged fish throughout the Willamette Basin above Willamette Falls, Oregon, by airplane and recorded their location. Fish migrated primarily during the spring to early summer period before stopping during the remainder of summer, when peak river temperatures (≥ 20°C) occurred. These fish tended to remain stationary through the fall and winter. However, at least a few fish continued to migrate upstream after September. I monitored maturation characteristics of adult Pacific lamprey, over time at Willamette Falls, Oregon and compared these fish with recent migrants collected from the Pacific Ocean as they entered freshwater. The results suggest a unimodal spawn timing between April and June, at water temperatures < 20 °C. Between July and mid-September, as water temperatures peaked at ~ 25 °C, relatively immature fish for both sexes prevailed. Warm summer temperatures coincided with an increase and prevalence of testicular atrophy in males, and I also observed a large die-off of lamprey during this time. The immature fish had maturation stages and phenotypic characteristics similar to recent migrants collected at the mouth of the Klamath River, suggesting that the immature fish at Willamette Falls would spawn the following year, and spawners in any given year may have been recent migrants during the previous year. However there is a temporal overlap in the spring of immature and mature fish, and I found evidence from gonad histology of maturing fish as they entered the river from the ocean, suggesting that a cohort is comprised of recent migrants that spawn within several weeks of entering freshwater, and another cohort is comprised of recent migrants that mature and spawn at least 1 year later. I hypothesize that the recent migrants that would likely spawn shortly after entering freshwater are akin to a winter or "ocean maturing" steelhead, Oncorhynchus mykiss, that optimizes feeding and growth in the open ocean for a few years before entering freshwater to spawn low in the river system shortly afterwards. Alternatively, these lamprey may be similar to coastal cutthroat trout, O. clarki clarki, that feed and grow in the coastal areas of the ocean for a few months before entering freshwater to spawn. There could be other less apparent explanations as well. I also hypothesize that the lamprey that would likely spawn within ~ 1 year of entering freshwater are akin to a "stream maturing" steelhead that foregoes feeding and growth opportunities, enters freshwater during the summer – fall, and accesses spawning grounds to spawn at temperatures that promote evolutionary fitness via successful spawning the following spring. Based on the results of my research, I hypothesize that warm summer temperatures (> 20 °C) can act as a strong selection factor against stream maturing Pacific lamprey in two ways. First, these temperatures may expedite their maturation, while at the same time slowing their migration. If these hypotheses are true, then I predict an uncoupling of spawn timing with optimal habitat characteristics, that would promote fitness, in the upper watershed. Second, summer temperatures may cause gonad atrophy and death prior to spawning. This scenario may select for ocean maturing Pacific lamprey. Title: The physiology ecology and run diversity of adult Pacific lamprey, Entosphenus tridentatus, during the freshwater spawning migration Author: Clemens, Benjamin Jacob, 1976- Pacific lamprey, Entosphenus tridentatus, have shown recent and rapid declines in abundance. These anadromous fish return to streams where they mature, spawn and die. It has been inferred that Pacific lamprey enter freshwater and reside for ~ 1 year before spawning. This long exposure to the freshwater environment may affect the plasticity of the maturation process and the migration timing of Pacific lamprey. Diversity in run times and body size has been observed for Pacific lamprey, yet it is unknown if this diversity is induced by the freshwater environment or if it is genetic. My first goal was to describe the maturation and migration characteristics of adult Pacific lamprey during their freshwater migration. My second goal was to use these data to make an estimation of the run diversity in Pacific lamprey. I conducted three complementary studies, in the laboratory and the field, to achieve these goals. I held immature adult lamprey (non-ripe fish that had ceased parasitic feeding in the ocean and had returned to freshwater) in the laboratory at temperatures that mimicked what these fish would experience in the wild, during the summer (mean: 21.8 °C), and another group of lamprey at cooler temperature (mean: 13.6 °C) to compare maturation timing and characteristics. The warm water group of lamprey showed significantly greater proportional decreases in body mass following temperature exposure than fish in the cooler water. All fish exposed to the warm water matured the following spring (8-10 months later) whereas only about half of the fish from the cool water exposure matured. To understand the migration distances and timing of adult Pacific lamprey, I tracked radio-tagged fish throughout the Willamette Basin above Willamette Falls, Oregon, by airplane and recorded their location. Fish migrated primarily during the spring to early summer period before stopping during the remainder of summer, when peak river temperatures (≥ 20°C) occurred. These fish tended to remain stationary through the fall and winter. However, at least a few fish continued to migrate upstream after September. I monitored maturation characteristics of adult Pacific lamprey, over time at Willamette Falls, Oregon and compared these fish with recent migrants collected from the Pacific Ocean as they entered freshwater. The results suggest a unimodal spawn timing between April and June, at water temperatures < 20 °C. Between July and mid-September, as water temperatures peaked at ~ 25 °C, relatively immature fish for both sexes prevailed. Warm summer temperatures coincided with an increase and prevalence of testicular atrophy in males, and I also observed a large die-off of lamprey during this time. The immature fish had maturation stages and phenotypic characteristics similar to recent migrants collected at the mouth of the Klamath River, suggesting that the immature fish at Willamette Falls would spawn the following year, and spawners in any given year may have been recent migrants during the previous year. However there is a temporal overlap in the spring of immature and mature fish, and I found evidence from gonad histology of maturing fish as they entered the river from the ocean, suggesting that a cohort is comprised of recent migrants that spawn within several weeks of entering freshwater, and another cohort is comprised of recent migrants that mature and spawn at least 1 year later. I hypothesize that the recent migrants that would likely spawn shortly after entering freshwater are akin to a winter or "ocean maturing" steelhead, Oncorhynchus mykiss, that optimizes feeding and growth in the open ocean for a few years before entering freshwater to spawn low in the river system shortly afterwards. Alternatively, these lamprey may be similar to coastal cutthroat trout, O. clarki clarki, that feed and grow in the coastal areas of the ocean for a few months before entering freshwater to spawn. There could be other less apparent explanations as well. I also hypothesize that the lamprey that would likely spawn within ~ 1 year of entering freshwater are akin to a "stream maturing" steelhead that foregoes feeding and growth opportunities, enters freshwater during the summer – fall, and accesses spawning grounds to spawn at temperatures that promote evolutionary fitness via successful spawning the following spring. Based on the results of my research, I hypothesize that warm summer temperatures (> 20 °C) can act as a strong selection factor against stream maturing Pacific lamprey in two ways. First, these temperatures may expedite their maturation, while at the same time slowing their migration. If these hypotheses are true, then I predict an uncoupling of spawn timing with optimal habitat characteristics, that would promote fitness, in the upper watershed. Second, summer temperatures may cause gonad atrophy and death prior to spawning. This scenario may select for ocean maturing Pacific lamprey. Close

• 2. [Article] The ocean ecology of salmon in the Northeast Pacific Ocean : an abridged history

  Research on the ecology of salmon in the northeast Pacific Ocean began in the early 20th century. Charles Gilbert and Willis Rich demonstrated the basis for the stock concept and were instrumental in changing ...

  Citation × Citation Citation Abstract Copy Title: The ocean ecology of salmon in the Northeast Pacific Ocean : an abridged history Author: McKinnell, Stewart M., Pearcy, William G. Research on the ecology of salmon in the northeast Pacific Ocean began in the early 20th century. Charles Gilbert and Willis Rich demonstrated the basis for the stock concept and were instrumental in changing common misconceptions of the times. Later in the 1900s, research endeavors, primarily under the auspices of the International North Pacific Fisheries Commission, led to important studies on the distribution and migration of maturing salmon on the high seas. Research on the early juvenile period was initiated later, especially after the 1982-1983 El Niño clearly showed the limits of the ocean's carrying capacity along the west coast of the United States. There is now good evidence for both intra- and interspecific competition among salmon in the open ocean and for correlations between variable physical environments, such as El Ninos and regime shifts, and survival of salmon during early ocean life. How mortality rates are affected by physical forcing, food availability, predation, and food web structure and how these effects will be modified by climate change and global warming are all major challenges for the future. Title: The ocean ecology of salmon in the Northeast Pacific Ocean : an abridged history Author: McKinnell, Stewart M., Pearcy, William G. Research on the ecology of salmon in the northeast Pacific Ocean began in the early 20th century. Charles Gilbert and Willis Rich demonstrated the basis for the stock concept and were instrumental in changing common misconceptions of the times. Later in the 1900s, research endeavors, primarily under the auspices of the International North Pacific Fisheries Commission, led to important studies on the distribution and migration of maturing salmon on the high seas. Research on the early juvenile period was initiated later, especially after the 1982-1983 El Niño clearly showed the limits of the ocean's carrying capacity along the west coast of the United States. There is now good evidence for both intra- and interspecific competition among salmon in the open ocean and for correlations between variable physical environments, such as El Ninos and regime shifts, and survival of salmon during early ocean life. How mortality rates are affected by physical forcing, food availability, predation, and food web structure and how these effects will be modified by climate change and global warming are all major challenges for the future. Close

• 3. [Article] Foraging Niche Separation of Adélie (Pygoscelis adeliae) and Gentoo (P. papua) Penguins During the Breeding Season at Palmer Station, Antarctica

  Climate-induced range overlap can result in novel interactions between similar species and potentially lead to competitive exclusion. The Western Antarctic Peninsula is one of the most rapidly warming ...

  Citation × Citation Citation Abstract Copy Title: Foraging Niche Separation of Adélie (Pygoscelis adeliae) and Gentoo (P. papua) Penguins During the Breeding Season at Palmer Station, Antarctica Author: Pickett, Erin Climate-induced range overlap can result in novel interactions between similar species and potentially lead to competitive exclusion. The Western Antarctic Peninsula is one of the most rapidly warming regions on Earth and is experiencing a poleward climate migration. This transition from a polar to sub-polar environment has resulted in a range expansion of the ice-intolerant gentoo penguin (Pygoscelis papua) and a coincident decline of ice-obligate Adélie penguins (P. adeliae) at Palmer Station, Anvers Island (64˚46’S, 64˚03’W). Ecologically similar species that share a limited prey resource must occupy disparate foraging niches in order to co-exist. Therefore, the aim of this study was to determine the extent of spatial and dietary niche segregation between Adélie and gentoo penguins during the breeding season at Palmer Station. This research was conducted across six austral breeding seasons, from 2010-2015, which allowed for an investigation of the consistency of niche overlap in the context of resource variability. This study was conducted using a twofold methodological approach involving biotelemetry and diet sampling, and little evidence was found to suggest that foraging competition is a primary driver of penguin population trajectories in this region. While substantial overlap was observed in the diets of Adélie and gentoo penguins, who consumed primarily Antarctic krill (Euphausia superba), these results show that Adélie and gentoo penguins partitioned this shared prey resource through horizontal and vertical segregation of their core foraging areas. No evidence was found to suggest that Antarctic krill is a limiting resource during the breeding season or that climate-induced sympatry of Adélie and gentoo penguins has resulted in competition for prey and the subsequent differing population trajectories. Rather, other physical and biological changes to the ecosystem due to the rapidly changing climate around the Antarctic Peninsula may have stronger influences on the current trends of these two species in this region. Title: Foraging Niche Separation of Adélie (Pygoscelis adeliae) and Gentoo (P. papua) Penguins During the Breeding Season at Palmer Station, Antarctica Author: Pickett, Erin Climate-induced range overlap can result in novel interactions between similar species and potentially lead to competitive exclusion. The Western Antarctic Peninsula is one of the most rapidly warming regions on Earth and is experiencing a poleward climate migration. This transition from a polar to sub-polar environment has resulted in a range expansion of the ice-intolerant gentoo penguin (Pygoscelis papua) and a coincident decline of ice-obligate Adélie penguins (P. adeliae) at Palmer Station, Anvers Island (64˚46’S, 64˚03’W). Ecologically similar species that share a limited prey resource must occupy disparate foraging niches in order to co-exist. Therefore, the aim of this study was to determine the extent of spatial and dietary niche segregation between Adélie and gentoo penguins during the breeding season at Palmer Station. This research was conducted across six austral breeding seasons, from 2010-2015, which allowed for an investigation of the consistency of niche overlap in the context of resource variability. This study was conducted using a twofold methodological approach involving biotelemetry and diet sampling, and little evidence was found to suggest that foraging competition is a primary driver of penguin population trajectories in this region. While substantial overlap was observed in the diets of Adélie and gentoo penguins, who consumed primarily Antarctic krill (Euphausia superba), these results show that Adélie and gentoo penguins partitioned this shared prey resource through horizontal and vertical segregation of their core foraging areas. No evidence was found to suggest that Antarctic krill is a limiting resource during the breeding season or that climate-induced sympatry of Adélie and gentoo penguins has resulted in competition for prey and the subsequent differing population trajectories. Rather, other physical and biological changes to the ecosystem due to the rapidly changing climate around the Antarctic Peninsula may have stronger influences on the current trends of these two species in this region. Close