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The Case Of The ‘Missing’ Arctic Bivalves And The Walrus: The Biggest [Overlooked] Clam Fishery On The Planet, Roger L. Mann, Eric N. Powell, Daphne M. Munroe
The Case Of The ‘Missing’ Arctic Bivalves And The Walrus: The Biggest [Overlooked] Clam Fishery On The Planet, Roger L. Mann, Eric N. Powell, Daphne M. Munroe
VIMS Articles
Bivalve molluscs represent a significant proportion of the diet of both Atlantic and Pacific walrus (Odobenus rosmarus rosmarus and Odobenus rosmarus divergens, respectively) and are pivotal to benthic–pelagic coupling and carbonate cycling in the Arctic oceans. The latter is of particular relevance in a period of seasonal ice retreat, freshwater release into associated surface waters, decreasing water pH, and possible undersaturation of Arctic waters with respect to aragonite. Using population estimates and predation rates for the walruses on bivalve molluscs, a conservative estimate of bivalve consumption in the regions of active walruses foraging is 2.0–3.0 3 106 tonnes y–1—a tonnage …
Estimating Shifts In Phenology And Habitat Use Of Cobia In Chesapeake Bay Under Climate Change, Daniel P. Crear, Brian E. Watkins, Marjorie A.M. Friedrichs, Pierre St-Laurent, Kevin C. Weng
Estimating Shifts In Phenology And Habitat Use Of Cobia In Chesapeake Bay Under Climate Change, Daniel P. Crear, Brian E. Watkins, Marjorie A.M. Friedrichs, Pierre St-Laurent, Kevin C. Weng
VIMS Articles
Cobia (Rachycentron canadum) is a large coastal pelagic fish species that represents an important fishery in many coastal Atlantic states of the U.S. They are heavily fished in Virginia when they migrate into Chesapeake Bay during the summer to spawn and feed. These coastal habitats have been subjected to warming and increased hypoxia which in turn could impact the timing of migration and the habitat suitability of Chesapeake Bay. With conditions expected to worsen, we project current and future habitat suitability of Chesapeake Bay for cobia and predict changes in their arrival and departure times as conditions shift. To do …
The Power Struggle: Assessing Interacting Global Change Stressors Via Experimental Studies On Sharks, Ian A. Bouyoucos, Sue-Ann Watson, Serge Planes, Colin A. Simpfendorfer, Gail D. Schwieterman, Et Al
The Power Struggle: Assessing Interacting Global Change Stressors Via Experimental Studies On Sharks, Ian A. Bouyoucos, Sue-Ann Watson, Serge Planes, Colin A. Simpfendorfer, Gail D. Schwieterman, Et Al
VIMS Articles
Ocean warming and acidification act concurrently on marine ectotherms with the potential for detrimental, synergistic effects; yet, effects of these stressors remain understudied in large predatory fishes, including sharks. We tested for behavioural and physiological responses of blacktip reef shark (Carcharhinus melanopterus) neonates to climate change relevant changes in temperature (28 and 31 °C) and carbon dioxide partial pressures (pCO2; 650 and 1050 µatm) using a fully factorial design. Behavioural assays (lateralisation, activity level) were conducted upon 7–13 days of acclimation, and physiological assays (hypoxia tolerance, oxygen uptake rates, acid–base and haematological status) were …
Sensitivity Of A Shark Nursery Habitat To A Changing Climate, Daniel Crear, Robert Latour, Marjorie A.M. Friedrichs, Pierre St-Laurent, Kevin C. Weng
Sensitivity Of A Shark Nursery Habitat To A Changing Climate, Daniel Crear, Robert Latour, Marjorie A.M. Friedrichs, Pierre St-Laurent, Kevin C. Weng
VIMS Articles
Nursery area habitats such as estuaries are vital for the success of many fish populations. Climate change is altering conditions in these areas, which can thus impact the availability of suitable nursery habitat. The sandbar shark Carcharhinus plumbeus uses Chesapeake Bay (USA) as a nursery habitat during the summer months from birth up to 10 yr of age. To assess the impacts of climate change on juvenile sandbar sharks, we developed a habitat model using longline data collected from a fishery-independent survey within Chesapeake Bay. With this model, we projected contemporary and future distributions of suitable habitat for juvenile sandbar …
Effects Of Seawater Exchange On Water Chemistry Among Coastal Lakes With Intermittent Connections To The Sea, A. Challen Hyman, Dana Bigham Stephens
Effects Of Seawater Exchange On Water Chemistry Among Coastal Lakes With Intermittent Connections To The Sea, A. Challen Hyman, Dana Bigham Stephens
VIMS Articles
Intermittently closed and open lakes and lagoons (ICOLLs) are a dynamic class of coastal waterbodies with the unique feature of intermittently connecting to the sea. Understanding the functioning and potential threats of these globally rare systems is important to their preservation and protection. Coastal dune lakes of northwest Florida are one example of an understudied group of ICOLLs which connect with the Gulf of Mexico for brief periods of time. Using a 17-year, monthly water chemistry dataset, we analyzed long-term patterns in water chemistry among 16 coastal dune lakes. Using salinity as a proxy for frequency of seawater inflows, principal …
Identifying Important Juvenile Dusky Shark Habitat In The Northwest Atlantic Ocean Using Acoustic Telemetry And Spatial Modeling, Charles W. Bangley, Tobey H. Curtis, David H. Secor, Robert J. Latour, Matthew B. Ogburn
Identifying Important Juvenile Dusky Shark Habitat In The Northwest Atlantic Ocean Using Acoustic Telemetry And Spatial Modeling, Charles W. Bangley, Tobey H. Curtis, David H. Secor, Robert J. Latour, Matthew B. Ogburn
VIMS Articles
Highly mobile species can be challenging for fisheries management and conservation due to large home ranges combined with dependence on discrete habitat areas where they can be easily targeted or vulnerable to anthropogenic disturbances. Management of the Dusky Shark Carcharhinus obscurus in the northwest Atlantic Ocean has been particularly challenging due to the species' inherent vulnerability to overfishing and poorly understood habitat associations. To better understand habitat associations and seasonal distributions, we combined telemetry and remotely sensed environmental data to spatially model juvenile Dusky Shark presence probability in the northwest Atlantic Ocean. To accomplish this, 22 juvenile Dusky Sharks (107-220 …
Spillover Of Sea Scallops From Rotational Closures In The Mid-Atlantic Bight (United States), Deborah R. Hart, Daphne M. Munroe, Joseph C. Caracappa, Dale Haidvogel, Burton V. Shank, David Rudders, Et Al
Spillover Of Sea Scallops From Rotational Closures In The Mid-Atlantic Bight (United States), Deborah R. Hart, Daphne M. Munroe, Joseph C. Caracappa, Dale Haidvogel, Burton V. Shank, David Rudders, Et Al
VIMS Articles
We examined evidence for larval spillover (increased recruitment outside the closures) of Atlantic sea scallops (Placopecten magellanicus) due to rotational closures in the Mid-Atlantic Bight using a 40-year fisheries survey time series and a larval transport model. Since the first closure of the Hudson Canyon South (HCS) area in 1998, mean recruitment in the two areas directly down-current from this closure, Elephant Trunk (ET) and Delmarva (DMV), increased significantly by factors of about 7 and 2, respectively. Stock–recruit plots indicate that low biomasses in HCS were associated with reduced mean recruitment in ET and DMV. Simulations indicate that …
Improving Production Of The Eastern Oyster Crassostrea Virginica Through Coordination Of Genetic Improvement Programs, Research, And Technology Transfer Activities, Standish K. Allen Jr., Caird Rexroad Iii, Robert Rheault
Improving Production Of The Eastern Oyster Crassostrea Virginica Through Coordination Of Genetic Improvement Programs, Research, And Technology Transfer Activities, Standish K. Allen Jr., Caird Rexroad Iii, Robert Rheault
VIMS Articles
The East Coast Shellfish Growers Association has been actively advocating for continued improvement of breeding infrastructure to support a vibrant and ecologically crucial industry. This white paper acknowledges these efforts and is meant to serve as a paradigm for oyster breeding on the East Coast. The vision is relatively straightforward; however, its execution—like most things—will be fraught with detail and inertia. Yet describing simply the idea without articulating some detail and potential roadblocks would be misleading. Instead, we present a picture of a breeding paradigm as it might play out across the entire range of a species. At this particular …
Evaluating Movements Of Opakapaka (Pristipomoides Filamentosus) Relative To A Restricted Fishing Area By Using Acoustic Telemetry And A Depth-Constrained Estimator Of Linear Home Ranges, Stephen R. Scherrer, Kevin C. Weng
Evaluating Movements Of Opakapaka (Pristipomoides Filamentosus) Relative To A Restricted Fishing Area By Using Acoustic Telemetry And A Depth-Constrained Estimator Of Linear Home Ranges, Stephen R. Scherrer, Kevin C. Weng
VIMS Articles
Networks of no-take fishery reserves have emerged as a tool for managing deepwater fish species. In Hawaii and elsewhere, such areas are used to manage deepwater snapper species. However, little is known regarding the movements of these species relative to protected areas. We used passive acoustic telemetry to track crimson jobfish (Pristipomoides filamentosus), also known as opakapaka, in one of Hawaii’s bottomfish restricted fishing areas to understand the size required for a reserve to protect this species. From January 2017 through January 2018, 179 fish were tagged. Only 10 fish were classified as alive on the basis of movements indicated …
Are Predator−Prey Model Predictions Supported By Empirical Data? Evidence For A Storm-Driven Shift Toan Alternative Stable State In A Crab−Clam System, Cassandra N. Glaspie, Rochelle D. Seitz, Romuald N. Lipcius
Are Predator−Prey Model Predictions Supported By Empirical Data? Evidence For A Storm-Driven Shift Toan Alternative Stable State In A Crab−Clam System, Cassandra N. Glaspie, Rochelle D. Seitz, Romuald N. Lipcius
VIMS Articles
A dynamic systems approach can predict steady states in predator−prey interactions,but there are very few examples of predictions from predator−prey models conforming to empirical data. Here, we examined the evidence for the low-density steady state predicted by a Lotka-Volterra model of a crab−clam predator−prey system using data from long-term monitoring, and data from a previously published field survey and field predation experiment. Changepoint analysis of time series data indicate that a shift to low density occurred for the soft-shell clam Mya arenaria in 1972, the year of Tropical Storm Agnes. A possible mechanism for the shift is that Agnes altered …
Fish Biodiversity Patterns Of A Mesophotic-To-Subphotic Artificial Reef Complex And Comparisons With Natural Substrates, St Jones, Jm Asher, Rc Boland, Bk Kanenaka, Kevin C. Weng
Fish Biodiversity Patterns Of A Mesophotic-To-Subphotic Artificial Reef Complex And Comparisons With Natural Substrates, St Jones, Jm Asher, Rc Boland, Bk Kanenaka, Kevin C. Weng
VIMS Articles
Artificial reefs act as high-rugosity habitats and are often deployed to enhance fishing; however, the effects of man-made features on fish communities can be unpredictable and are poorly understood in deeper waters. In this study, we used a submersible to describe a deep-water artificial reef complex (93-245 m) off of Ewa Beach, Oahu, Hawaii, USA, and evaluated possible conservation and/or fisheries-related contributions. Sixty-eight species were recorded, with larger features supporting greater diversity of species. Species composition changed strongly with depth and a faunal break was detected from 113-137 m. While the features supported diverse fish communities, they were not similar …
Bycatch In A Commercial Lobster Fishery: Effects On Two Benthic Predators, Sea Raven And Longhorn Sculpin, Brooke N. Anderson, Amelia M. Weissman, Brett Sweezey, John Mandelman, David Rudders, James A. Sulikowski
Bycatch In A Commercial Lobster Fishery: Effects On Two Benthic Predators, Sea Raven And Longhorn Sculpin, Brooke N. Anderson, Amelia M. Weissman, Brett Sweezey, John Mandelman, David Rudders, James A. Sulikowski
VIMS Articles
Studying the species‐specific responses to fishing capture is critical for effective management and conservation of bycatch species given that acute stress incurred from capture and handling may ultimately lead to mortality. While species of low commercial value are often overlooked, having accurate information on the effects of capture on all species is necessary for ecosystem‐based management. Sea Raven (SR) Hemitripterus americanus and Longhorn Sculpin (LHS) Myoxocephalus octodecemspinosus are routinely captured in the commercial American lobster Homarus americanus fishery in the Gulf of Maine, and they are discarded due to low commercial value. Despite a lack of economic value, these predatory …
In The Face Of Climate Change And Exhaustive Exercise: The Physiological Response Of An Important Recreational Fish Species, Daniel P. Crear, Richard Brill, Lauren M.L. Averilla, Sara C. Meakem, Kevin C. Weng
In The Face Of Climate Change And Exhaustive Exercise: The Physiological Response Of An Important Recreational Fish Species, Daniel P. Crear, Richard Brill, Lauren M.L. Averilla, Sara C. Meakem, Kevin C. Weng
VIMS Articles
Cobia (Rachycentron canadum) support recreational fisheries along the US mid- and south-Atlantic states and have been recently subjected to increased fishing effort, primarily during their spawning season in coastal habitats where increasing temperatures and expanding hypoxic zones are occurring due to climate change. We therefore undertook a study to quantify the physiological abilities of cobia to withstand increases in temperature and hypoxia, including their ability to recover from exhaustive exercise. Respirometry was conducted on cobia from Chesapeake Bay to determine aerobic scope, critical oxygen saturation, ventilation volume and the time to recover from exhaustive exercise under temperature and …
Barriers To Eastern Oyster Aquaculture Expansion In Virginia, Jennifer Beckensteiner, David M. Kaplan, Andrew M. Scheld
Barriers To Eastern Oyster Aquaculture Expansion In Virginia, Jennifer Beckensteiner, David M. Kaplan, Andrew M. Scheld
VIMS Articles
The eastern oyster once provided major societal and ecosystem benefits, but these benefits have been threatened in recent decades by large declines in oyster harvests. In many areas, recovery of oyster aquaculture faces significant societal opposition and spatial constraints limiting its ability to meet expectations regarding future food needs and provision of ecosystem services. In Virginia, oyster aquaculture has begun to expand, concurrent with an increase in subaqueous leased areas (over 130,000 acres of grounds are currently leased). Though private leases must in theory be used for oyster production, in practice, they can be held for other reasons, such as …
Timing Of The Reproductive Cycle Of Waved Whelk, Buccinum Undatum, On The Us Mid-Atlantic Bight, Sarah Borsetti, Daphne Munroe, David Rudders, Jul-Han Chang
Timing Of The Reproductive Cycle Of Waved Whelk, Buccinum Undatum, On The Us Mid-Atlantic Bight, Sarah Borsetti, Daphne Munroe, David Rudders, Jul-Han Chang
VIMS Articles
Development of the unmanaged waved whelk (Buccinum undatum) fishery on the Mid-Atlantic continental shelf of the United States has initiated investigation into fisheries-related biological and population attributes of the species in this region. Maturation and reproduction timing vary by location for this species and are likely linked to bottom water temperature. This study examined the seasonal fluctuations in relevant body metrics and gonadosomatic index in relation to bottom temperature to assess the timing of the reproductive cycle of the B. undatum population in the southern-most extent of this species' range in the Atlantic. To characterize variation over the maturation schedule, …
Density-Dependence Mediates The Effects Of Temperature On Growth Of Juvenile Blue Catfish In Nonnative Habitats, Vaskar Nepal, Mary C. Fabrizio
Density-Dependence Mediates The Effects Of Temperature On Growth Of Juvenile Blue Catfish In Nonnative Habitats, Vaskar Nepal, Mary C. Fabrizio
VIMS Articles
The combined effects of conspecific density and climate warming on the vital rates of invasive fish species have not been well studied, but may be important in predicting how successful they will be in the future. We evaluated the effects of temperature and population density on monthly time series of sizes of age-0 Blue Catfish Ictalurus furcatus in the James, York, and Rappahannock River subestuaries (defined here as tidally influenced bodies of water that feed into the Chesapeake Bay) from 1996 to 2017, using growing degree-days (GDDs, °C day) as a measure of thermal time. Our pre- dictive linear mixed-effects …
Movements, Habitat Utilization, And Post-Release Survival Of Cobia (Rachycentron Canadum) That Summer In Virginia Waters Assessed Using Pop-Up Satellite Archival Tags, Douglas R. Jensen, John Graves
Movements, Habitat Utilization, And Post-Release Survival Of Cobia (Rachycentron Canadum) That Summer In Virginia Waters Assessed Using Pop-Up Satellite Archival Tags, Douglas R. Jensen, John Graves
VIMS Articles
Cobia (Rachycentron canadum) is a cosmopolitan marine fish that inhabits tropical, sub-tropical, and temperate marine and estuarine waters and supports a major recreational fishery along the U.S. Atlantic and Gulf of Mexico coasts. Recent changes in U.S. cobia management have sparked controversy and highlighted limitations in our understanding of the species’ biology. This study utilized pop-up satellite archival tags (PSATs) to assess the movements, habitat utilization, and post-release survival of cobia that summer in Virginia waters
Investigating The Efficacy Of A Proposed Marine Protected Area For The Endangered Humphead Wrasse Cheilinus Undulatus At A Remote Island Group In Seychelles, Ryan Daly, Clare A. Keating, Andrew E. Gray, Lauren R. Peel, Luke Gordon, James S.E. Lea, Christopher R. Clarke, Kevin C. Weng
Investigating The Efficacy Of A Proposed Marine Protected Area For The Endangered Humphead Wrasse Cheilinus Undulatus At A Remote Island Group In Seychelles, Ryan Daly, Clare A. Keating, Andrew E. Gray, Lauren R. Peel, Luke Gordon, James S.E. Lea, Christopher R. Clarke, Kevin C. Weng
VIMS Articles
The humphead wrasse Cheilinus undulatus is an iconic, ecologically important and Endangered fish species associated with coral reefs in the Indo-Pacific region. Due to its large size and complex life history characteristics, it is vulnerable to overfishing and has undergone substantial population declines in parts of its range. Knowledge of the species’ movement ecology is currently limited to only 2 previous studies, and very little is known about populations in the western Indian Ocean. The present study aimed to use passive acoustic telemetry to investigate the importance of a remote coral reef to a population of humphead wrasse in the …
Drivers And Trends In Catch Of Benthic Resources In Chilean Turfs And Surrounding Open Access Areas, Jennifer Beckensteiner, Andrew M. Scheld, Miriam Fernandez, David M. Kaplan
Drivers And Trends In Catch Of Benthic Resources In Chilean Turfs And Surrounding Open Access Areas, Jennifer Beckensteiner, Andrew M. Scheld, Miriam Fernandez, David M. Kaplan
VIMS Articles
Beginning in the 1990’s, Chile implemented an extensive Territorial User Rights for Fisheries (TURFs) network that now comprises nearly 1,000 TURFs. This network provides a rare opportunity to examine spatial and temporal trends in TURF use and impacts on surrounding open access areas (OAAs). In this analysis, landings of keyhole limpet (Fissurella spp.), kelp (Lessonia spp.) and red sea urchin (Loxechinus albus) were used to estimate catch-per-unit effort (CPUEs) and catch-per-unit area (CPUAs) indices inside and outside TURFs by fishing cove. For these species, CPUEs and CPUAs in 2015 were significantly higher inside TURFs. However, temporal trends analyzed with a …
Contemporary And Future Distributions Of Cobia, Rachycentron Canadum, Dp Crear, Be Watkins, Vs Saba, Je Graves, Dr Jensen, Aj Hobday, Kevin C. Weng
Contemporary And Future Distributions Of Cobia, Rachycentron Canadum, Dp Crear, Be Watkins, Vs Saba, Je Graves, Dr Jensen, Aj Hobday, Kevin C. Weng
VIMS Articles
Climate change has influenced the distribution and phenology of marine species, globally. However, knowledge of the impacts of climate change is lacking for many species that support valuable recreational fisheries. Cobia (Rachycentron canadum) are the target of an important recreational fishery along the U.S. east coast that is currently the subject of a management controversy regarding allocation and stock structure. Further, the current and probable future distributions of this migratory species are unclear, further complicating decision-making. The objectives of this study are to better define the contemporary distribution of cobia along the U.S. east coast and to project potential shifts …
Sublethal Effects Of Salinity And Temperature On Non-Native Blue Catfish: Implications For Establishment In Atlantic Slope Drainages, Vaskar Nepal, Mary C. Fabrizio
Sublethal Effects Of Salinity And Temperature On Non-Native Blue Catfish: Implications For Establishment In Atlantic Slope Drainages, Vaskar Nepal, Mary C. Fabrizio
VIMS Articles
The distribution and further range expansion of non-native blue catfish Ictalurus furcatus in coastal waters throughout the United States Atlantic slope depend, in part, on the salinity tolerance of the fish. However, temperature-mediated sublethal effects of increased salinities on blue catfish biology are not yet known. We assessed the effects of salinity and temperature on growth, body condition, body composition and food consumption of juvenile blue catfish in a controlled laboratory experiment. Temperature and salinity had an interactive effect on blue catfish biology, although most fish survived 112 days in salinities up to 10 psu. At salinities 9 psu) likely …
Quantifying The Behavioral And Economic Effects Of Regulatory Change In A Recreational Cobia Fishery, Andrew M. Scheld, William M. Goldsmith, Shelby White, Hamish J. Small, Susanna Musick
Quantifying The Behavioral And Economic Effects Of Regulatory Change In A Recreational Cobia Fishery, Andrew M. Scheld, William M. Goldsmith, Shelby White, Hamish J. Small, Susanna Musick
VIMS Articles
Fisheries economists typically assume recreational anglers make decisions that maximize individual angler utility, which may depend on fishery and regulatory conditions. Under this framework, changes in regulations can lead to target species substitution by anglers in response to shifts in expectations of trip utility. A stated preference survey was developed and distributed to recreational cobia (Rachycentron canadum) anglers in Virginia to explore the effects of regulatory change on angler decision-making, species targeting, and resulting economic outcomes. The survey included a series of hypothetical choice scenarios, where respondents were asked to select their most preferred alternative after being presented with different …
Bony-Tongue Fishes (Teleostei: Osteoglossomorpha) From The Eocene Nanjemoy Formation, Virginia, Eric J. Hilton, Jeffrey Carpenter
Bony-Tongue Fishes (Teleostei: Osteoglossomorpha) From The Eocene Nanjemoy Formation, Virginia, Eric J. Hilton, Jeffrey Carpenter
VIMS Articles
Bony-tongue fishes, Osteoglossomorpha, are distributed in North and South America, Africa, Asia, and Australia and are found on all continents except Antarctica in the fossil record. The group includes fishes such as the mooneyes (Hiodontidae), freshwater knifefishes (Notopteridae), elephantfishes (Mormyridae), and the arowanas and pirarucu (Osteoglossidae). Remains identified as belonging to the family Osteoglossidae are known from the Nanjemoy Formation of Maryland and northern Virginia and comprise isolated teeth and fragmentary jaw bones assigned to the now extinct †Brychaetus muelleri. The second author discovereda partial toothed parasphenoid among other isolated and frag-mentary vertebrate microfossils from the Fisher–Sullivan Site of the …
Comparative Transcriptomics Of Spotted Seatrout (Cynoscion Nebulosus) Populations To Cold And Heat Stress, Jingwei Song, Jan Mcdowell
Comparative Transcriptomics Of Spotted Seatrout (Cynoscion Nebulosus) Populations To Cold And Heat Stress, Jingwei Song, Jan Mcdowell
VIMS Articles
Resilience to climate change depends on a species' adaptive potential and phenotypic plasticity. The latter can enhance survival of individual organisms during short periods of extreme environmental perturbations, allowing genetic adaptation to take place over generations. Along the U.S. East Coast, estuarine‐dependent spotted seatrout (Cynoscion nebulosus) populations span a steep temperature gradient that provides an ideal opportunity to explore the molecular basis of phenotypic plasticity. Genetically distinct spotted seatrout sampled from a northern and a southern population were exposed to acute cold and heat stress (5 biological replicates in each treatment and control group), and their transcriptomic responses …
Invasive Blue Catfish In The Chesapeake Bay Region: A Case Study Of Competing Management Objectives, Mary C. Fabrizio, Vaskar Nepal, Troy D. Tuckey
Invasive Blue Catfish In The Chesapeake Bay Region: A Case Study Of Competing Management Objectives, Mary C. Fabrizio, Vaskar Nepal, Troy D. Tuckey
VIMS Articles
Freshwater fishes have been introduced outside their native range to establish recreational fisheries, but management conflicts arise when such introductions also result in potentially harmful effects on native species. In this case study, we focus on Blue Catfish Ictalurus furcatus, which were introduced in the Chesapeake Bay region and are now considered invasive. In many tidal tributaries, Blue Catfish have increased dramatically in abundance, expanded into high-salinity habitats (up to 21.8 psu), and negatively affected native species, prompting calls for the development of an effective management plan. However, management of this conflict species is complicated by multiple competing objectives, including …