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    Assessment of the Yellowstone Lake food web during lake trout suppression and Yellowstone cutthroat trout recovery informs conservation benchmarks
    (Montana State University - Bozeman, College of Letters & Science, 2022) Glassic, Hayley Corrine; Chairperson, Graduate Committee: Christopher S. Guy; This is a manuscript style paper that includes co-authored chapters.
    The collapse of native Yellowstone cutthroat trout Oncorhynchus clarkii bouvieri (hereafter cutthroat trout) in Yellowstone Lake was caused by predation by invasive lake trout Salvelinus namaycush. As an ecosystem with a low-diversity fish assemblage and several longterm data sets, Yellowstone Lake provided a unique opportunity to evaluate the influence of an invasive salmonid population undergoing suppression beyond only predator-prey dynamics. Diet data for cutthroat trout and lake trout were evaluated at varying densities to determine the effects of density on diet composition and diet plasticity. During the lake trout high-density state, lake trout consumed fewer native cutthroat trout and switched to amphipods, which were also consumed by cutthroat trout, resulting in high diet overlap between the species. As suppression reduced invasive lake trout densities, lake trout returned to consuming cutthroat trout and diet overlap was released. A shift in lake trout delta 13C signatures from the high-density state to the moderate-density state also corroborates higher consumption of cutthroat trout and invasive lake trout diet plasticity. Beyond predator-prey dynamics of lake trout and cutthroat trout, the invasion of lake trout caused > or = 25% change in energy flux for all organisms in Yellowstone Lake except for copepods. Food-web functional state did not change among food webs, but percentage of functional state contributing to total flux did vary. Herbivory was the dominant food-web functional state for all years, with the greatest percentage of flux from herbivory in 2011. In addition, by using a whole-ecosystem model that accounted for whirling disease and historical (natural) lake-level variation, I show that suppression of the lake trout population is necessary for cutthroat trout recovery, but the amount of suppression effort needed for cutthroat trout to reach recovery benchmarks is linked to severity of climate change. Additionally, if climate change increases the frequency and severity of reduced lake levels in the future, cutthroat trout recovery benchmarks may need to be adapted. With this research, I demonstrate how the feedbacks among predator-prey dynamics, disease, and climate change can complicate the suppression of invasive species and the conservation of invaded ecosystems and must be considered for establishing realistic conservation benchmarks.
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    Combination of acoustic telemetry and side-scan sonar provides insight for lake trout Salvelinus namaycush suppression in a submontane lake
    (Montana State University - Bozeman, College of Letters & Science, 2021) Siemiantkowski, Michael James; Chairperson, Graduate Committee: Christopher S. Guy
    Expansion of an invasive Lake Trout Salvelinus namaycush population in Swan Lake, Montana threatens a core area population of Bull Trout Salvelinus confluentus in Montana. Given the increased efficacy of suppression using novel embryo suppression methods, there is renewed interest in Lake Trout suppression in Swan Lake. The specific questions of this study were: 1) where are Lake Trout spawning, 2) where are the most used spawning sites, 3) what is the amount of spawning habitat, 4) does the estimated spawning area differ between estimates from telemetry locations and side-scan sonar imagery of suitable spawning substrate, and 5) how much phosphorous and nitrogen would be added to Swan Lake if carcass-analog pellet treatments were implemented? Acoustic tags were implanted in 85 Lake Trout in July and August of 2018 and 2019. Nightly tracking efforts during September, October, and November of 2018 and 2019 resulted in 1,744 relocations for 49 individual Lake Trout. Kernel-density analysis was used to evaluate Lake Trout aggregation locations identifying 10 distinct spawning sites -- corroborating previous studies. Visual observation of Lake Trout embryos confirmed spawning at three sites with the remaining seven sites considered to be unconfirmed spawning sites. All confirmed spawning sites were located in the littoral zone along areas of steep bathymetric relief and were the most used across both spawning seasons. In 2019, side-scan sonar imaging was used to classify and quantify the total area of suitable spawning substrate, which comprised 12.8% of the total surface area estimated for confirmed sites and 11.4% for unconfirmed spawning sites. Simultaneous treatment of all confirmed and unconfirmed spawning sites would require 205,709 + or - 86 kg of carcass-analog pellet material, resulting in 370.4 + or - 0.2 kg of phosphorous and 7,487.9 + or - 3.1 kg of nitrogen inputs to Swan Lake. Thus, pellet treatment would increase the Carlson's trophic state index (TSI) values from 20.8 to 27.7 for total phosphorous, and from 22.1 to 26.2 for total nitrogen. Based on a TSI threshold value of < 40 for an oligotrophic lake, the use of carcass-analog pellets could be a feasible addition to renewed Lake Trout suppression efforts in Swan Lake.
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    Non-target effects of a novel invasive species management strategy: benthic invertebrate responses to lake trout embryo suppression in Yellowstone Lake, Wyoming
    (Montana State University - Bozeman, College of Letters & Science, 2020) Briggs, Michelle Anne; Chairperson, Graduate Committee: Lindsey Albertson; Lindsey K. Albertson, Dominique R. Lujan, Lusha M. Tronstad, Hayley C. Glassic, Christopher S. Guy and Todd M. Koel were co-authors of the article, 'Carcassd deposition to suppress invasive lake trout causes differential mortality of two common benthic invertebrates in Yellowstone Lake, Wyoming' in the journal 'Fundamental and applied limnology' which is contained within this thesis.; Lindsey K. Albertson, Dominique R. Lujan, Lusha M. Tronstad, Hayley C. Glassic, Christopher S. Guy and Todd M. Koel were co-authors of the article, 'Non-target effects of a novel suppression technique for invasive fishes: responses of benthic invertebrate communities' submitted to the journal 'Ecological applications' which is contained within this thesis.
    Invasive species threaten native biodiversity and ecosystem function, and suppression is often required to reduce these effects. However, invasive species management actions can cause harmful, unintended consequences for non-target taxa. In Yellowstone Lake, Wyoming, invasive lake trout (Salvelinus namaycush) have reduced abundance of the native Yellowstone cutthroat trout (Oncorhynchus clarkii bouvieri), decreasing availability of an important food source for aquatic and terrestrial predators. Gillnets are used to suppress adult lake trout, and the lake trout carcasses are then deposited onto spawning sites in the littoral zone to cause embryo mortality by reducing dissolved oxygen concentrations as they decay. However, this management action may have non-target effects on organisms in the lake, including benthic invertebrates, which comprise a large portion of native trout diets. Some taxa of invertebrates may benefit from the addition of nutrients to the littoral zone, while other taxa may experience mortality in response to low dissolved oxygen conditions caused by carcass decay. We conducted two field experiments to understand how carcass treatment affects benthic invertebrates in Yellowstone Lake. First, we conducted an in situ experiment with individual invertebrates housed in small chambers covered by carcasses to determine if carcass treatment causes mortality of hypoxia-tolerant amphipods and hypoxia-sensitive caddisflies. We found that carcass treatment caused increased mortality in caddisflies but not amphipods. Second, we conducted a field experiment to investigate how carcass treatment affects invertebrate communities when applied at entire spawning sites. We also compared invertebrate communities at cobble-dominated lake trout spawning sites to macrophyte-dominated sites to determine if carcass treatment could alter food web dynamics at a lake-wide scale. We found that carcass treatment causes non-target effects on benthic invertebrates, specifically reducing immobile taxa, hypoxia-sensitive taxa, and Chironomidae, and altering community structure. Areas dominated by macrophytes had more abundant and larger invertebrates than spawning sites. Due to the small spatial extent of spawning sites and the higher abundance of invertebrates at other habitats in the lake, we conclude carcass treatment can have localized non-target effects at a local scale but is unlikely to alter food-web dynamics at a lake-wide scale.
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    Evaluation of embryo suppression methods for nonnative lake trout in Yellowstone Lake, Yellowstone National Park, Wyoming, USA
    (Montana State University - Bozeman, College of Letters & Science, 2019) Poole, Alex Stephen; Chairperson, Graduate Committee: Alexander V. Zale
    Introduced Lake Trout Salvelinus namaycush threaten native Yellowstone Cutthroat Trout Oncorhynchus clarkii bouvieri in Yellowstone Lake, Yellowstone National Park. Gill nets have been used to suppress subadult and adult Lake Trout since 1995. Because survival of embryonic and larval life history stages can have profound effects on population dynamics of Lake Trout, suppression at those stages, especially if used in concert with intensive gill netting of older fish, could enhance suppression efforts. Therefore, I conducted controlled laboratory and field experiments to systematically evaluate the effects of a variety of candidate chemical (sodium chloride, calcium carbonate, gelatin, and liquid and powdered rotenone), biological (carcass and carcass analog), and physical (sediment) suppression methods on different developmental stages of Lake Trout embryos and larvae. Liquid and powdered rotenone applications, fish carcass and carcass analog exposures, and sediment deposition significantly increased embryo mortality in laboratory experiments. Sodium chloride, calcium carbonate, and gelatin applications were not effective. In-situ exposure to ground carcass material in Yellowstone Lake resulted in 100% embryo mortality in 14 and 28 kg/m 2 biomass treatments; sediment deposition caused 97% embryo mortality among overwintering incubators. Embryo mortality was probably caused by hypoxic conditions within substrates. Embryo suppression methods differed in their effectiveness, rate at which mortality was achieved, and ease of application. These differences, as well as Lake Trout spawning site characteristics such as depth, contour, fetch, substrate size, interstitial depth, isolation, and presence of non-target organisms ultimately determine which embryo suppression method will be most applicable in a given situation. Nevertheless, implementation of successful embryo suppression techniques evaluated in this study could be used to increase mortality of Lake Trout in Yellowstone Lake. Incorporating effective embryo suppression in an Integrated Pest Management approach has the potential to provide more effective Lake Trout suppression in the long term.
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    Quantifying the spatial structure of invasive lake trout in Yellowstone Lake to improve suppression efficacy
    (Montana State University - Bozeman, College of Letters & Science, 2019) Williams, Jacob Robert; Chairperson, Graduate Committee: Christopher S. Guy
    Conserving Yellowstone Cutthroat Trout by suppressing invasive Lake Trout in Yellowstone Lake is a high priority for Yellowstone National Park natural-resource managers. Insight into the spatial structure of Lake Trout throughout the lake will help increase the efficacy of the Lake Trout suppression program. Lake Trout (N = 578) were surgically implanted with dual acoustic and radio transmitters from 2015 through 2017. Mobile acoustic (boat) and radio (fixed-wing aircraft) telemetry surveys were performed to identify aggregations of Lake Trout. Telemetry surveys occurred during the spawning period (autumn) in 2016 and during the summer and spawning period in 2017. Lake Trout exhibited distinct aggregations during the summer and spawning period. Lake Trout aggregated at nine locations during the summer 2017 and were most frequently located in the West Thumb. Lake Trout aggregated at 22 locations during the spawning period including 12 previously undocumented putative spawning locations. Two aggregations in the West Thumb, Carrington Island and Anglers Bluff, had the highest relative densities of Lake Trout. Aggregations during the summer were generally farther from shore, greater in depth, and more dispersed than aggregations during the spawning period. Targeting locations of Lake Trout, as identified through telemetry, with gill nets was an effective strategy for increasing catch-per-unit-effort. The Lake Trout suppression program is probably altering the behavior of Lake Trout in Yellowstone Lake, which explains the high number of spawning locations and low spawning site fidelity relative to other research studies on Lake Trout spawning behavior. This study provided valuable insight into the spatial structure of Lake Trout in Yellowstone Lake. The areas Lake Trout aggregated will continue to be targeted by gillnetting and novel embryo suppression methods.
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    Evaluation of suppression methods targeting non-native lake trout embryos in Yellowstone Lake, Yellowstone National Park, Wyoming, USA
    (Montana State University - Bozeman, College of Letters & Science, 2017) Thomas, Nathan Andrew; Chairperson, Graduate Committee: Christopher S. Guy
    Non-native Lake Trout Salvelinus namaycush threaten to extirpate native Yellowstone Cutthroat Trout Oncorhynchus clarkii bouvieri in Yellowstone Lake, Yellowstone National Park. Suppression of Lake Trout in Yellowstone Lake has been ongoing since 1995, primarily by gillnetting. Unfortunately, bycatch of Yellowstone Cutthroat Trout is associated with this removal method, which targets adult and subadult Lake Trout. Alternative methods effective at causing mortality in Lake Trout early life stage(s) could be used simultaneously with gillnetting to improve suppression effectiveness. The vulnerability of salmonid embryos suggest increasing Lake Trout embryo mortality is feasible and because population growth rates are sensitive to age-0 survival an effective embryo suppression method could have population-level effects. Thus, the primary objective of this study was to evaluate the efficacy of methods to increase mortality of Lake Trout embryos. In situ experiments tested the effect of suction dredging, electroshocking, tarping, and Lake Trout carcass deposition on embryo mortality. The secondary objective of this study was to evaluate interstitial water flow using NaCl dry injection to better understand the feasibility of using chemicals for embryo suppression. Concurrent laboratory studies have shown that Lake Trout embryos experience high mortality after chemical exposure. Tarping and suction dredging were not effective at increasing embryo mortality. Electroshocking caused 0.92 mortality of embryos at the substrate surface but only 0.38 at 20 cm depth in the substrate. Lake Trout carcass deposition caused 0.99 mortality of embryos, both at the surface and at 20 cm in the substrate. The direction and rate of interstitial water flow was not consistent and future work on a larger scale is needed to inform the feasibility of chemical suppression. Hypoxic conditions within the carcass sites may have caused embryo mortality. In addition, carbon dioxide CO 2 and hydrogen sulfide H 2S are byproducts of organic matter decomposition that harm developing salmonid embryos. Embryo suppression methods are unlikely to replace traditional Lake Trout suppression methods. However, the success of Lake Trout carcass deposition shows potential for the development of an effective additional suppression method that could be implemented on a large scale.
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    Use of otolith microchemistry to identify Yellowstone cutthroat trout and lake trout natal origins and movement patterns in Yellowstone Lake, Wyoming
    (Montana State University - Bozeman, College of Letters & Science, 2016) Stewart, Kole Patrick; Chairperson, Graduate Committee: Thomas E. McMahon
    The Yellowstone Lake Yellowstone cutthroat trout population has declined as a result of drought, whirling disease, and the introduction of lake trout. Little is known about the recruitment patterns of cutthroat trout and lake trout in this system. Otolith microchemistry is uniquely suited for answering these questions by matching the chemical signatures found in otoliths to the same signatures found in the water fish occupy. My first objective was to identify and compare the primary spawning streams contributing to historic (1997) and recent (2013) cutthroat trout recruitment. I analyzed the chemical signatures (87 Sr:86 Sr, Sr:Ca, Ba:Ca, Mg:Ca, and Mn:Ca) of 22 cutthroat trout spawning streams and the same signatures from the natal region of cutthroat trout otoliths. There was low variation among the chemical signatures of many spawning streams, thus streams were grouped into 9 clusters using a cluster analysis. Relative recruitment to each cluster was assessed using random forest models with a classification accuracy of 84.4% for known-origin cutthroat trout fry otoliths and 79.0% for simulated otolith signatures. There was a significant difference in the proportions of recruitment between historic and recent cutthroat trout spawning clusters (X 2 = 15.40, p = 0.03). The majority of historic (0.84) and recent (0.77) recruitment occurred in the same three stream clusters, with the most notable change being a decrease in recent recruitment in the stream cluster containing Pelican Creek and an increase in recruitment in tributaries in the upper Yellowstone River drainage. The second objective was to identify the spawning locations and movement patterns of lake trout within Yellowstone Lake. I analyzed the 87 Sr:86 Sr, and Sr:Ca signatures from 8 locations throughout Yellowstone Lake and the same signatures in 20 lake trout otoliths. I did not find sufficient variation within the lake water chemistry to differentiate lake regions and there was no significant differences found within in the lake trout otolith transects. This study can be used to inform future spawning stream conservation and restoration by directing managers towards spawning streams of increasing or decreasing importance. This study also highlights some of the strengths and limitations of using microchemistry studies in freshwater system.
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    Dynamics of Yellowstone cutthroat trout and lake trout in the Yellowstone Lake ecosystem : a case study for the ecology and management of non-native fishes
    (Montana State University - Bozeman, College of Letters & Science, 2015) Syslo, John Michael; Chairperson, Graduate Committee: Christopher S. Guy; Christopher S. Guy and Todd M. Koel were co-authors of the article, 'Trophic overlap and temporal diet shifts for a nonnative and a native salmonid in Yellowstone Lake, Yellowstone National Park' submitted to the journal 'Transactions of the American Fisheries Society' which is contained within this thesis.; Christopher S. Guy, Todd M. Koel, Patricia E. Bigelow, Philip D. Doepke, Brian D. Ertel and Jeffrey L. Arnold were co-authors of the article, 'Response of Yellowstone cutthroat trout to suppression of non-native lake trout in the Yellowstone Lake ecosystem' submitted to the journal 'Canadian journal of fisheries and aquatic sciences' which is contained within this thesis.
    The introduction of lake trout Salvelinus namaycush into Yellowstone Lake preceded the collapse of the native Yellowstone cutthroat trout Oncorhynchus clarkii bouvieri population. As a system with a simple fish assemblage and several long-term data sets, Yellowstone Lake provided a unique opportunity to evaluate the ecology of a native salmonid in the presence of a non-native salmonid population undergoing suppression in a large natural lake. Diet data for Yellowstone cutthroat trout and lake trout were evaluated at varying densities to determine the effects of density on diet composition. Temporal diet shifts from 1996-1999 to 2011-2013 were likely caused by limitation of prey fish for lake trout. Diets, stable isotopes, and depth-related patterns in CPUE indicated lake trout > 300 mm consumed primarily amphipods, making them trophically similar to Yellowstone cutthroat trout from during 2011-2013. A lake trout removal program was initiated during 1995 to reduce predation on Yellowstone cutthroat trout. Abundance and fishing mortality were estimated for lake trout from 1998 through 2013 and Yellowstone cutthroat trout from 1986 through 2013. Density-dependence was evaluated by examining individual growth, weight, maturity, and pre-recruit survival as a function of abundance. In addition, a simulation model was developed for the lake trout- Yellowstone cutthroat trout system to determine the probability of Yellowstone cutthroat trout abundance persisting at performance metrics given potential reductions in lake trout abundance. Estimates of Yellowstone cutthroat trout abundance varied 5-fold and lake trout abundance varied 6-fold. Yellowstone cutthroat trout weight and pre-recruit survival decreased with increasing Yellowstone cutthroat trout abundance; however, individual growth and maturity were not related to abundance. Lake trout population metrics did not vary with lake trout abundance. Simulation model results were variable because of uncertainty in lake trout pre-recruit survival. Conservative estimates for required lake trout reductions were > 97% of 2013 abundance for a > 70% probability of Yellowstone cutthroat trout persistence at the performance metrics outlined in the Native Fish Conservation Plan. Lake trout removal will likely reduce lake trout abundance and result in Yellowstone cutthroat trout recovery if the amount of fishing effort exerted in 2013 is maintained for at least 15 years.
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    Efficacy of suppressing non-native lake trout in an isolated backcountry lake in Glacier National Park
    (Montana State University - Bozeman, College of Letters & Science, 2014) Fredenberg, Carter Roger; Chairperson, Graduate Committee: Christopher S. Guy
    Prior to the recent invasion of non-native lake trout Salvelinus namaycush, Glacier National Park (GNP) supported approximately one-third of the remaining natural lake habitat supporting threatened bull trout Salvelinus confluentus. However, bull trout populations have recently declined and are at high risk of extirpation in several lakes in western GNP due to the establishment of lake trout. In 2009, the U.S. Geological Survey and the National Park Service began suppressing lake trout in Quartz Lake (352 ha) to reduce impacts to native bull trout. The objectives of this study were to: (1) describe the demography of the lake trout population during the suppression program (2009-2013); (2) identify the timing and location of lake trout spawning; (3) determine the most efficient combination of gill net mesh color and twine diameter to capture juvenile lake trout (age 2 to age 4); (4) assess the effects of suppression on the growth rate of the lake trout population and use this information to model harvest scenarios; and (5) determine whether suppression negatively impacted bull trout. Lake trout exhibited slower growth, lower condition, and lower fecundity relative to other populations. Spawning locations were identified on cobble and boulder substrates (depths 2-20 m) near the base of two avalanche chutes where adults began aggregating between 1 and 9 October prior to thermal destratification (11-12 C°). Catch rates of spawning (ripe) adults were highest 12-25 October when temperatures declined to below 10 C°. Gill nets with 0.1 mm twine thickness and green color increased catchability of juvenile lake trout. Although density dependent parameters were not included, population simulation models indicated the population was growing exponentially and would likely reach carrying capacity within ten years without suppression. Suppression resulted in declining population growth rates (lambda) from 1.23 prior to suppression to 0.61-0.79 during suppression. Bull trout redd abundances remained stable throughout the suppression period. My results indicate targeted suppression successfully reduced lake trout abundance and that continued suppression at or above observed exploitation levels is needed to ensure continued population declines and to avoid impacts to the bull trout population.
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    Relative contributions of climate variation, lake trout predation, and other factors to the decline of Yellowstone Lake cutthroat trout during the three recent decades
    (Montana State University - Bozeman, College of Letters & Science, 2010) Kaeding, Lynn Robert; Chairperson, Graduate Committee: Daniel Goodman; Thomas E. McMahon (co-chair)
    The relative contributions of climate variation, lake trout Salvelinus namaycush predation, and other factors to the recent, three-decade decline of the lacustrine-adfluvial (i.e., a life-history form consisting of fish that mostly live in a lake but spawn in an inflowing tributary) Yellowstone cutthroat trout Oncorhynchus clarkii bouvieri (YCT) population of Clear Creek, a Yellowstone Lake tributary, were evaluated. Strong growth of that population's storied spawning run between the early 1960s and 1978, when the run peaked at about 70,000 fish, had been considered key evidence of recovery of the lake's YCT population from formerly excessive angler harvest and other adverse factors. Thus the run's subsequent, almost continuous decline to about 500 fish in 2007 was perplexing. Gillnet catches of YCT at established lake locations likewise indicated a concurrent decline in the lake-wide YCT population. Prominent among the factors that may have importantly affected the YCT population during the recent decades was predation by the illegally introduced, reproducing, nonnative lake trout discovered in Yellowstone Lake in 1994. Data mainly taken from YCT in the spawning run (n = 29 years) and gillnet catch (n = 30 years) were examined for information useful to specifying the Leslie matrix of a dynamic, age-structured model that had climate as a covariate. The model, fitted to spawning run size and mean total length (TL) of YCT in the run during 1977-2007 (n = 29 data years), explained 87% of variation in observed run size, 86% of variation in observed mean TL, and strongly suggested that climate (as indexed by total-annual air degree-days > 0°C measured on the lake's north shore) had an important effect on recruitment of age-0 YCT to subsequent spawning runs. Results also suggested that an effect of lake trout predation on survival of age-1 to age-5 YCT became apparent only during the recent decade. The important test of ongoing efforts to control lake trout in Yellowstone Lake and thereby limit their predation on YCT - on the basis of data for YCT - will occur when climatic conditions improve for YCT recruitment to the Clear Creek and other YCT spawning stocks of the lake.
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