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Carbon dioxide (CO2) gas and eDNA monitoring as tools for eradicating invasive fish from anchialine pools in Hawai‘i

Invasive fish can profoundly affect communities they invade. In Hawai‘i, invasive fishes have become established in many anchialine pools, threatening the persistence of resident invertebrates, including several endangered species. Tools to eradicate invasive fishes from these pools are lacking. This study tested the efficacy of carbon dioxide (CO2) gas diffused into anchialine pool water as a method to eradicate invasive Mozambique tilapia (Oreochromis mossambicus), guppies (Poecilia reticulata), and western mosquitofish (Gambusia affinis). We first conducted aquarium trials to identify how these fishes were affected by elevated CO2 and the concomitant reduction in pH. We then carried out field trials in pools containing these fish in one pool each at two national historical parks on the Island of Hawai‘i during July 2021–January 2022. We also developed environmental DNA (eDNA) protocols to detect fish that may have survived CO2 treatments. The effect of CO2 on fish behavior varied among species; at pH 5.3 (CO2 = 255 mg/L) for tilapia and 5.0 (CO2 = 488 mg/L) for tilapia, guppies, and mosquitofish, all generally lost their ability to swim, showed slow or no gill movement, and altered their position in the water column. No tilapia survived the trials (n = 4 and 6 individuals at pH 5.3 and 5.0, respectively). In contrast, 41.7% (n = 12) of adult guppies and 66.7% (n = 12) of adult mosquitofish survived treatment at pH 5.0. In the field we were unable to reduce anchialine pool water pH below 5.7. Regardless, we were able to eradicate tilapia from one pool over four sequential treatments. Post-treatment eDNA assessments supported visual surveys, confirming our results. We were not able to eradicate guppies and mosquitofish. Results from this study show that CO2 can be an effective tool for eradicating invasive tilapia from anchialine pools, and post-treatment eDNA assessments can provide managers with a method for evaluating the success of eradication efforts.

Hawaii

Identifying research in support of the management and control of dreissenid mussels in the western United States

On February 9–10, 2022, the Pacific States Marine Fisheries Commission, U.S. Fish and Wildlife Service, U.S. Geological Survey, and Washington State University hosted a workshop to establish research priorities that support the implementation of action items listed in a current invasive species management plan, the Quagga and Zebra Mussel Action Plan (QZAP) 2.0, that are intended to limit the establishment and spread of quagga and zebra mussels in the western United States. The workshop focus was on developing research priorities for the thematic areas that are addressed in QZAP 2.0: 1) early detection monitoring, 2) prevention and containment, 3) control and management, and 4) rapid response. In addition, research priorities were developed for a fifth thematic area that addressed dreissenid mussel biology. Forty scientists participated in the two-day workshop. Prior to the workshop, participants were asked to review and rank research priorities that were established for a previous version of the QZAP and to offer suggestions on emerging research priorities. During the workshop, subject matter experts presented information describing current knowledge of research and information associated with the thematic areas of early detection monitoring, prevention and containment, rapid response, control and management, and biology in the context of strategies and actions listed in QZAP 2.0. The rankings of previous research priorities and suggestions of emerging priorities were then reviewed, and a revised list of research priorities was formed. The list of research priorities is presented by thematic area.

Management of Biological Invasions

Experimental observations on the decay of environmental DNA from bighead and silver carps

Interest in the field of environmental DNA (eDNA) is growing rapidly and eDNA surveys are becoming an important consideration for aquatic resource managers dealing with invasive species. However, in order for eDNA monitoring to mature as a research and management tool, there are several critical knowledge gaps that must be filled. One such gap is the fate of eDNA materials in the aquatic environment. Understanding the environmental factors that influence the decay of eDNA and how these factors impact detection probabilities over time and space could have significant implications for eDNA survey design and data interpretation. Here we experimentally explore decay of eDNA associated with bighead carp ( Hypophthalmichthys nobilis ) biological waste collected from an aquaculture filtration system and with sperm collected from captive silver carp ( H. molitrix ), and how decay may be influenced by differing levels of water turbulence, temperature, microbial load, and pH. We found that the decay patterns of eDNA associated with both H. nobilis biological waste and H. molitrix milt significantly fit monophasic exponential decay curves. Secondly, we observed that the highest temperature we tested resulted in a decay half-life as much as 5.5× more rapid than the lowest temperature we tested. When we suppressed microbial loads in eDNA samples, we observed that overall losses of eDNA were reduced by about 2.5×. When we amended eDNA samples with pond water the half-life of eDNA was reduced by about 2.25×, despite relatively little apparent increase in the overall microbial load. This pattern indicated that species constituency of the microbial community, in addition to microbial load, might play a critical role in eDNA degradation. A shift in pH from 6.5 to 8.0 in the samples resulted in a 1.6× reduction in eDNA halflife. Water turbulence in our study had no apparent effect on eDNA decay. When we combined different temperature, pH, and microbial load treatments to create a rapid decay condition and a slow decay condition, and tracked eDNA decay over 91 days, we observed a 5.0× greater loss of eDNA by Day 5 under rapid decay conditions than under slow decay conditions. At the end of the trials, the differences in eDNA loss between the rapid decay and baseline and slow decay conditions were 0.1× and 3.3×, respectively. Our results strongly demonstrate the potential for environmental factors to influence eDNA fate and, thus, the interpretation of eDNA survey results.

Management of Biological Invasions

America's most wanted fishes: Cataloging risk assessments to prioritize invasive species for management action

Hundreds of fish species enter the United States through human intervention (e.g., importation) and some of these fishes pose a substantial risk to the nation’s assets and ecosystems. Prevention, early detection, and rapid response (EDRR) are vital to stop species invasions, but time and resources to manage the large suite of fish species that enter the nation are limited. Evaluating the risk of a species’ invasion in a location is one way to prioritize among many species for management action. Species risk assessments are often associated with information systems or are published within grey literature or peer-reviewed journal articles. Improving access to available risk assessments could help in prioritizing management action for the most potentially invasive fish species. We aggregated fish species risk assessments, synthesizing the current knowledge on the risk of fish invasions in the United States. To accomplish this, we searched information systems and conducted a literature review. We then summarized risk assessment results along with the importation statuses of fish species and identified if imported, high-risk species are managed under federal or state policy. Within the scope of the conterminous U.S., we found 98 high-risk fish species. Eighteen of these species are imported to the country, but only three species have been recently prohibited from importation according to the Lacey Act. We observed similar patterns at the scales of the Great Lakes region and Florida. Collectively, our work provides a baseline estimate of the high-risk fish invaders that enter the U.S. through importation, underscoring species to consider for priority management action, as well as a benchmark of species that lack risk assessments. Insights from this work can be enriched when joined with other invasive species information, which could be accomplished through a national EDRR information system, an information sharing hub in development by the U.S. Geological Survey.

Management of Biological Invasions

Assessing grass carp (Ctenopharyngodon idella) occupancy and detection probability within Lake Erie from environmental DNA

Grass carp ( Ctenopharyngodon idella ), an invasive cyprinid within the Laurentian Great Lakes, is naturally reproducing in several Lake Erie tributaries, which has raised concerns of the species’ spread throughout Lake Erie and the other Great Lakes. Knowledge of the recent invasion extent outside of the western basin of Lake Erie, particularly in eastern tributaries and nearshore waters, is limited. Understanding the invasion extent would improve the efficacy of ongoing coordinated multi-agency control efforts. Molecular tools, such as environmental DNA (eDNA), have shown promise for early detection of aquatic invasive species. In this study, water samples (N = 476) were collected for grass carp eDNA monthly between May and November in 2018 and 2019, at three sites in the Michigan waters of Lake Erie and the Detroit River. We fit Bayesian multi-scale occupancy models to determine differences in eDNA capture and detection probability among grass carp qPCR assays, sampling sites, and across time. To determine whether grass carp were physically present, and to validate eDNA samples, we quantified recent grass carp presence in sampled areas using an existing acoustic telemetry and field sampling framework. Our results indicate that grass carp eDNA capture probability differed among sites, but there was no difference among months. Positive grass carp eDNA detections were observed across multiple months at each site, with 69% of site-specific sampling events testing positive for grass carp eDNA on at least one assay and replicate. The majority (65%) of weeks where positive eDNA sampling detections occurred also concurrently had one or more grass carp detected via acoustic telemetry 1–6 days prior. Our results highlight the potential utility of using eDNA to monitor the invasion extent of grass carp within the nearshore waters of Lake Erie. However, further evaluation of the factors that influence grass carp eDNA characteristics among sites within Lake Erie are needed to determine its efficacy for surveillance protocols by natural resource management agencies.

Lake Erie

Evaluating the piscicide rotenone as an option for eradication of invasive Mozambique tilapia in a Hawaiian brackish-water wetland complex

Mozambique tilapia Oreochromis mossambicus were recently discovered in ‘Aimakapā Fishpond, a 12-hectare brackish-water wetland complex in Kaloko-Honokōhau National Historical Park, on the Island of Hawai’i. As a possible eradication method, we evaluated rotenone, a natural piscicide used in fish management and the active ingredient in plants traditionally used by indigenous Hawaiians for capturing fish. To assess rotenone’s efficacy in killing tilapia and effects on non-target species, laboratory toxicity tests involved exposing organisms to various concentrations of liquid CFT Legumine (5% rotenone) in static trials of 48-h to 72-h duration. Test organisms included: Mozambique tilapia, non-native guppy Poecilia reticulata , the non-native odonate Rambur’s forktail Ischnura ramburii , native feeble shrimp Palaemon debilis , and native ‘ōpae’ula shrimp Halocaridina rubra . All organisms and water used in tests were obtained from ‘Aimakapā (12.6–12.7 ppt salinity), or, for H. rubra , an anchialine pool (15.0–15.2 ppt salinity). Survival analyses indicated CFT Legumine concentrations >3 ppm (>0.15 mg/L rotenone) achieved 100% mortality of tilapia and 93% of guppies within 24 h, with most tilapia killed by 6 h and most guppies by 2 h. Little or no mortality was observed among invertebrate exposed to 1 to 5 mg/L CFT Legumine: 0% mortality for ‘ōpae’ula shrimp, 4% for feeble shrimp; and 16% for odonate larvae. The 48 h LC50 values for Mozambique tilapia and guppy were 0.06 and 0.11 mg/L rotenone, respectively. Results demonstrate rotenone’s potential for non-native fish eradication in brackish-water habitats, with benefit of low mortality to certain macro-invertebrates. High rotenone tolerance displayed by ‘ōpae’ula shrimp is noteworthy. Invasive fish are common in anchialine pools, threatening existence of shrimp and other invertebrate fauna. Although rotenone’s effects on freshwater organisms have been well studied, our research represents one of only a few controlled laboratory experiments quantitatively assessing rotenone tolerance of brackish or marine fauna.

Hawaii

What it is to be established: policy and management implications for non-native and invasive species

Management of invasive species, whether prevention, population reduction, or eradication, requires assessment of the invasive species’ population status and an assessment of the probability of success of management options. Perceptions of a species’ permanence in an environment or lack thereof frequently drives how limited time, financial, and personnel resources are allocated to such efforts. Language we use to describe a non-native species’ status largely defines these perceptions and sets boundaries, real or perceived, to potential management actions. Here we discuss the use of a particular term – “established” – when confronting management decisions for invasive species. Our objective is to contribute to bridging the gap between the realms of conceptual development and management with respect to use of the term “established”. We find that although there are benefits of polysemy and synonymy to conceptual development they present an additional challenge to managers who must weigh the costs, benefits, and potential for success of particular management actions. We also examine how existing conceptual frameworks might be augmented to bridge the theoretical-practical gap, such as more precisely defining potential management actions and explicitly including assessment of risk.

Management of Biological Invasions

Using structured decision making to evaluate potential management responses to detection of dreissenid mussel (Dreissena spp.) environmental DNA

Environmental (e)DNA tools are sensitive and cost-effective for early detection of invasive species. However, the uncertainty associated with the interpretation of positive eDNA detections makes it challenging to determine appropriate natural resource management responses. Multiple sources of error can give rise to positive detections of eDNA in a sample when individuals of that species are not present at the site or a widespread infestation is not imminent. Acting on an erroneous eDNA inference could result in needless costs or reductions in desirable resources. Alternatively, failure to rapidly act on eDNA results that truly indicate invader presence could compound negative impacts and lead to high, long-term costs to manage infestations. We used a structured decision making (SDM) process, which incorporates tradeoffs and uncertainties, to evaluate appropriate response actions following hypothetical eDNA detections of invasive dreissenid mussel ( Dreissena spp.) eDNA in Jordanelle Reservoir, Utah (USA). We worked with decision-makers and stakeholders to identify objectives and discrete management action alternatives to assess consequences and tradeoffs. Alternatives ranged from no action to intensive and expensive control efforts. The best performing alternative was delayed containment described by immediate attempts to confirm the eDNA detections using non-molecular sampling techniques followed by mandatory watercraft exit inspections to prevent dreissenid mussel spread to regional water bodies. Non-molecular sampling increased public support for management by demonstrating a commitment to monitor the invasion state before action, whereas containment decreased likelihood of regional spread to other waters. Delayed containment had the lowest downside risk, and the highest upside gains relative to other alternative actions. Sensitivity analyses showed our results to be robust to parameter and outcome uncertainty.

Management of Biological Invasions

Integrating landscape simulation models with economic and decision tools for invasive species control

In managing invasive species, land managers and policy makers need information to help allocate scarce resources as efficiently and effectively as possible. Decisions regarding treatment methods, locations, effort, and timing can be informed by the integration of landscape simulation models with economic tools. State and transition simulation models align with conceptual models of ecosystem change often used by practitioners and have been used to characterize the potential consequences of invasions. Outputs of these simulations are typically summarized to describe landscape changes (e.g., infested area), which may provide sufficient information for mangers to make informed decisions. However, it is sometimes helpful or necessary to go a step further to consider the social and economic values associated with treating (or not treating) invasions. Here, we describe when and how to integrate state and transition simulation models with economic and decision tools to aid in the control of emerging and established populations of invasive species. The paper provides an overview of three types of questions that can be addressed: 1) how big is the problem? 2) which management strategy is most appropriate? and 3) what are key sources of uncertainty? For each question, we describe aspects that can be addressed by landscape simulation models alone, and outstanding questions that can be evaluated by integrating economic and decision tools. Through a series of example applications from the literature, we reinforce how the integration of these tools, and the interdisciplinary perspective such an integration requires, can increase relevance and utility of modeling efforts for resource managers and decision makers.

Management of Biological Invasions

A workshop to advance invasive species early detection capacity of The Rapid Environmental DNA Assessment and Deployment Initiative & Network (READI-Net)

Early detection and rapid response (EDRR) can minimize the impacts of invasive species, which cost billions of dollars globally. To bolster EDRR across the United States, the U.S. Department of the Interior is working with the U.S. Geological Survey and other partners to advance a National EDRR Framework that strengthens tools, actions, and processes to find and eradicate invasive species before they establish and cause negative impacts. An important component of this framework is to strengthen molecular tools for detecting new invasions. The Rapid Environmental (e)DNA Assessment and Deployment Initiative & Network (READI-Net) project is developing automating eDNA sampling tools, processes to ensure that eDNA results are reliable for management decision-making, and information structures to deliver eDNA results to end-users. To improve the potential uptake of this molecular EDRR toolbox, READI-Net investigators met with a group of end-users, partners, developers and subject-matter experts from federal agencies, tribes, universities, and an NGO representing state agencies from February 28 to March 1, 2023, in Moss Landing, CA. Here, we summarize the READI-Net project and the corresponding participant feedback.

Management of Biological Invasions

Pathways of fish invasions in the Mid-Atlantic region of the United States

Non-native fish introductions are a major threat to biodiversity and fisheries, and occur through numerous pathways that vary regionally in importance. A key strategy for managing invasions is to focus prevention efforts on pathways posing the greatest risk of future introductions. We identified high-risk pathways for fish establishment in the Mid-Atlantic region of the United States based on estimates of probability of establishment and records of previous introductions, which were considered in the context of emerging socioeconomic trends. We used estimates of propagule pressure, species’ environmental tolerance, and size of species pool to assess the risk of establishment by pathway. Pathways varied considerably in historic importance and species composition, with the majority of species introduced intentionally via stocking (primarily for sport, forage, or biocontrol) or bait release. Bait release, private stocking, illegal introductions intended to establish reproducing populations (e.g., of sport fish), aquaculture, and the sale of live organisms all create risks for future invasions in the Mid-Atlantic region. Of these pathways, bait release probably poses the greatest risk of introductions for the Mid-Atlantic region because propagule pressure is moderate, most released species are tolerant of local environmental conditions, and the pool of species available for transplantation is large. Our findings differ considerably from studies in other regions (e.g., bait release is a dominant pathway in the Mid-Atlantic region, whereas illegal introduction of sport fish is dominant in the western US and aquarium releases are dominant in Florida), demonstrating the need for regional-scale assessments of, and management strategies for, introduction pathways.

Delaware, Maryland, New York, North Carolina, Penn

READI-Net—Providing tools for the early detection and management of aquatic invasive species

Overview Early detection of biological threats, such as invasive species, increases the likelihood that control efforts will be successful and cost-effective. Environmental deoxyribonucleic acid (eDNA) sampling is an established method for the efficient and sensitive early detection of new biological threats. The Rapid eDNA Assessment and Deployment Initiative & Network (READI-Net) is a project designed with partners to provide a full suite of tools to maximize the power of eDNA sampling for detecting invasive species. The READI-Net suite of tools will include the availability of autonomous eDNA samplers, multispecies molecular DNA detection tools, strategic sample design, standardized and repeatable lab analysis, and a communication framework to deliver eDNA detection results to inform invasive species science, policy, and management. The READI-Net project is part of a national strategy to implement eDNA sampling for the early detection of and rapid response to biological threats.

Fact Sheet

Ontogenetic and ecological variation in invasion risk of Brown Treesnakes (Boiga irregularis) on Guam

Size structure within populations of invasive species may have consequences for relative risk at all stages of the invasion process, with implications for management interventions such as interdiction, suppression, and eradication. To assess relative distributions of invasive Brown Treesnakes among demographic categories of management interest, we undertook the most comprehensive and controlled sampling in >25 years of research into this ecologically and economically destructive introduced predator. We collected a seasonally-balanced sample of 100 snakes from each of 18 sites, stratified by six habitat types, encompassing the species’ entire extralimital range. Samples indicated significant differences in distributions of female and male snakes among management classes (juvenile, transitional, and mature) by site and habitat. We found substantial heterogeneity in localized population characteristics over relatively small geographic distances, only modest influence of habitat type, higher prevalence of reproductively mature snakes in savanna and urban habitats, and an alarmingly high proportion of snakes that are too small to be effectively targeted by current rodent-baited control tools (mean = 38.2%, range = 19 to 72%). Failure to account for such variability in high risk demographic fractions may hinder successful interventions.

Guam

Survival and behavior of Chinese mystery snails (Bellamya chinensis) in response to simulated water body drawdowns and extended air exposure

Nonnative invasive mollusks degrade aquatic ecosystems and induce economic losses worldwide. Extended air exposure through water body drawdown is one management action used for control. In North America, the Chinese mystery snail (Bellamya chinensis) is an invasive aquatic snail with an expanding range, but eradication methods for this species are not well documented. We assessed the ability of B. chinensis to survive different durations of air exposure, and observed behavioral responses prior to, during, and following desiccation events. Individual B. chinensis specimens survived air exposure in a laboratory setting for > 9 weeks, and survivorship was greater among adults than juveniles. Several B. chinensis specimens responded to desiccation by sealing their opercula and/or burrowing in mud substrate. Our results indicate that drawdowns alone may not be an effective means of eliminating B. chinensis. This study lays the groundwork for future management research that may determine the effectiveness of drawdowns when combined with factors such as extreme temperatures, predation, or molluscicides.

Management of Biological Invasions

Ligation and division of ductus deferens does not produce long term sterility in most bighead carp or grass carp

Invading species are most easily eradicated or controlled if detected early and rapid action can be taken, but locating and eradicating small numbers of aquatic invaders is extremely difficult. Bighead carp ( Hypophthalmichthys nobilis ) and grass carp ( Ctenopharyngodon idella ) are Asian cyprinids that have been widely introduced and are considered undesirable and detrimental invasive species in many parts of their introduced range. The relatively novel “Judas fish” technique is used to locate aggregations of fish in the wild by strategically releasing individuals equipped with surgically implanted transmitters. However, releasing Judas fish that are capable of reproducing is problematic because it could add to the invasion problem by contributing to recruitment if not recaptured. We thus tested whether surgical ligation and division of the ductus deferens could effectively sterilize adult diploid bighead and grass carp. If effective, surgical sterilization would result in a superior Judas fish by maintaining the reproductive motivation derived from intact gonads while blocking the reproductive potential. This technique was initially successful, but ultimately most individuals were able to recanalize the surgically-severed ducts and produce milt, and were thus potentially able to reproduce during the following spawning season. Thus, ligation and division of the ductus deferens does not seem to be a viable procedure to sterilize bighead carp or grass carp for deployment as Judas fish where long term sterility of the fish is paramount in importance.

Missouri

Comparative olfactory and behavioral response of grass carp ( Ctenopharyngodon idella ), bighead carp ( Hypophthalmichthys nobilis ), silver carp ( H. molitrix ) and black carp ( Mylopharyngodon piceus ) to chemical stimuli

Grass carp ( Ctenopharyngodon idella ), bighead carp ( Hypophthalmichthys nobilis ), silver carp (H. molitrix), and black carp ( Mylopharyngodon piceus ), collectively “invasive carp,” represent an ongoing management problem in the Mississippi and Missouri River basins of North America. Development of strategies to control their populations are also ongoing. As a follow-up to previous research, we tested two amino acids and an amino acid-based commercial feeding stimulant as olfactory stimulants that may attract or repel these species. We employed a two-pronged approach to test how invasive carp respond to the olfactory stimulants, using an electro-olfactogram (EOG) to measure olfactory response and a tank with a stimulant in one half to test behavioral response. Consistent with previous research, we found grass carp had a higher magnitude of EOG response to stimulants compared to bighead and, marginally, silver carp. Bighead, silver, and black carp did not differ from each other in EOG response. Grass carp produced significantly greater lysine and a marginally greater alanine EOG response than bighead and silver carp and significantly greater commercial feeding stimulant response than bighead carp. Black carp had marginally greater lysine response than bighead carp. In the behavioral tests, grass carp generally avoided the stimuli tested, while bighead carp were generally attracted to stimuli. Neither black nor silver carp showed a clear attraction or avoidance toward the stimuli tested. To further evaluate these solutions for management applications additional testing in ponds or open water may be necessary.

Management of Biological Invasions

Tradeoffs of a portable, field-based environmental DNA platform for detecting invasive northern pike (Esox lucius) in Alaska

Environmental DNA (eDNA) has improved detection probabilities of aquatic invasive species but lab-based analytical platforms for eDNA analyses slow opportunities for rapid response. Effective approaches that address this analytical bottleneck and improve capacity for rapid response are urgently needed. We tested the sensitivity of a portable, field-based eDNA platform relative to widely used lab-based eDNA approaches for detecting invasive northern pike (Esox lucius) in eight lakes on Alaska’s Kenai Peninsula. The portable, field-based platform takes ~ 1 hr from sample collection to final results and uses a field-based DNA extraction kit, a shelf-stable assay, and a portable real-time PCR thermocycler. Lab-based approaches take days to weeks to months for final results and use lab-based DNA extraction kits, lab-bound assays, and benchtop real-time thermocyclers. We found that the portable, field-based approach was less sensitive than lab-based approaches and was more prone to inhibition, thus increasing potential for false-negatives. Until sensitivity and inhibition issues can be resolved, this portable, field-based approach is best viewed as a complement to rather than a replacement of standard eDNA lab-based approaches.

Alaska

Toxicity of copper sulfate and rotenone to Chinese mystery snail ( Bellamya chinensis )

The Chinese mystery snail ( Bellamya chinensis ) is a freshwater snail native to Southeast Asia, Japan, and Russia and is currently classified as an invasive species in at least 27 states in the USA. The species tolerates a wide range of environmental conditions, making management of established populations difficult. We tested the efficacy of two traditional chemical treatments, rotenone and copper sulfate, on the elimination of adult Chinese mystery snails in laboratory experiments. All snails (N=50) survived 72-hour exposure to rotenone-treated lake water, and 96% (N=25) survived 72-hour exposure to pre-determined rotenone concentrations of 0.25, 2.5, and 25.0 mg/L. All snails (N=10) survived exposure to 1.25 mg/L copper sulfate solution, 90% (N=10) survived exposure to 2.50 mg/L copper sulfate solution, and 80% (N=5) survived exposure to 5.0 mg/L copper sulfate solution. Neither rotenone nor copper sulfate effectively killed adult Chinese mystery snails in laboratory experiments, most likely due to their relatively large size, thick shell, and operculum. Therefore, it appears that populations will be very difficult to control once established, and management should focus on preventing additional spread or introductions of this species.

Management of Biological Invasions