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At least 253 records · Page 14Linked to original sources
USGS2021

Larval Coregonus spp. diets and zooplankton community patterns in the Apostle Islands, Lake Superior

With the exception of lake whitefish ( Coregonus clupeaformis ), relatively little is known about the early life history of larval coregonines in the Laurentian Great Lakes. For example, our knowledge of the feeding ecology of larval coregonines (excluding lake whitefish) is based on only 900 stomachs reported in the literature. Here, we describe the diets and demographics of larval coregonines from ice-out to late July, and the contemporaneous zooplankton community, in the Apostle Islands region of Lake Superior in 2018. Exogenous feeding was evident among the smallest larvae (down to 6 mm). Percent of larvae with food in their stomachs increased and yolk reserves decreased as larvae grew from 10 to 13 mm. A majority of the diet (58%) was copepod nauplii, with generally positive selection for adult copepods and Holopedium . The patterns in exogenous feeding and yolk sac absorption were similar to observations in Lake Superior in the 1970s. Diets were also generally similar, although Limnocalanus , Holopedium , and zooplankton eggs were more prevalent in 2018 than the 1970s. Demographic data suggested at least two distinct cohorts and/or coregonine species in 2018. Post-hoc genetic testing of larvae in a parallel study suggested our samples comprised a mix of predominantly cisco ( C. artedi ), kiyi ( C. kiyi ), and bloater ( C. hoyi ). Early life history studies, when coupled with emerging genetic techniques that can identify larval coregonines to species, will provide a powerful combination to better understand population dynamics of coregonines at a time of ongoing restoration and rehabilitation efforts throughout the Great Lakes.

Apostle Islands, Lake Superior↗
USGS2021

Assessing the assumptions of classification agreement, accuracy, and predictable healing time of sea lamprey wounds on lake trout

Sea lamprey control in the Laurentian Great Lakes relies on records of sea lamprey wounds on lake trout to assess whether control efforts are supporting fisheries management targets. Wounding records have been maintained for 70 years under the assumption that they are a reliable and accurate reflection of sea lamprey damage inflicted on fish populations. However, two key assumptions underpinning the use of these data need thorough evaluation: sea lamprey wounds follow a predictable healing progression, and individuals classify wounds accurately and reliably. To assess these assumptions, we conducted a workshop where experienced professionals examined lake trout with known sea lamprey wounds. For most lake trout, pictures were taken at regular intervals during the healing process. Our evaluation of wound pictures found high variability in healing times and wound progressions that did not conform to the currently used classification system. Participants’ wound classification agreement and accuracy were low and misclassification rates were high for most wound types. Training provided during the workshops did not markedly improve these metrics. We assessed wound classification accuracy for the first time and found assumptions of high accuracy and agreement are not met. We recommend misclassification rates be incorporated into models using wound data, sensitivity analyses be conducted to assess the potential impact of wound misclassification on estimates of key metrics (such as sea lamprey-induced mortality for lake trout), and alternative biomarkers be developed to quantify wound status with greater accuracy and precision.

Laurentian Great Lakes↗
USGS2020

Managing native and non-native sea lamprey ( Petromyzon marinus ) through anthropogenic change: A prospective assessment of key threats and uncertainties

Sea lamprey ( Petromyzon marinus ) is a species of conservation concern in their native range of the Atlantic coasts of Europe (Near Threatened to Critically Endangered) and North America (Secure to Critically Imperiled), and an invasive species of great economic and ecological concern in the Laurentian Great Lakes. Despite differences in life history strategy (anadromous natives vs adfluvial non-natives), the biology of sea lamprey is sufficiently similar to expect comparable responses to large-scale environmental change. We take a prospective look at the future (50 to 100 years) of sea lamprey management in an era of considerable environmental disturbance, and consider biological responses, management actions, and the future status of populations across the native and non-native ranges. Based on facilitated discussion by a diverse group of international experts, two major but poorly characterized classes of threats to sea lamprey were identified: climate change and socio-political issues. We discuss how climate induced changes affect growth, bioenergetics, and phenology of sea lamprey, and associated effects on control tactics (pesticides and barriers) and conservation. We consider tensions surrounding improving connectivity in the Great Lakes while controlling invasive sea lamprey, and discuss supplements and alternatives to pesticides and their wider effect, as well as the effects of new invasive species. To prevent the extirpation of native sea lamprey populations, or the re-expansion of non-native populations, we conclude with a call for new and ongoing dialogue and collaboration among all sea lamprey biologists and managers across the native and non-native range.

Journal of Great Lakes Research↗
USGS2020

Determining habitat limitations of Maumee River walleye production to western Lake Erie fish stocks: Documenting a spawning ground barrier

Tributaries provide spawning habitat for three of four major sub-stocks of Lake Erie walleye ( Sander vitreus ). Despite anthropogenic degradation and the extirpation of other potamodromous species, the Maumee River, Ohio, USA continues to support one of the largest fish migrations in the Laurentian Great Lakes. To determine if spawning habitat availability and quality could limit production of Maumee River walleye, two habitat suitability models were created for the lower 51 km of the Maumee River and the distribution and numbers of walleye eggs deposited in a 25 km stretch of river were assessed. Walleye eggs were collected using a diaphragm pump at 7 and 10 sites from March/April to May 2014 and 2015. The habitat suitability models showed that <3% of the river yielded ‘good’ walleye spawning habitat and 11–38% yielded ‘moderate’ walleye spawning habitat, depending on the model. However, a large set of rapids at river kilometer 28 and more than five river kilometers of less suitable habitat separated areas of ‘good’ habitat. The rapids may present a migratory barrier for many spawning walleye, as modeled water velocities exceed maximum estimated walleye swim speeds 71–100% of days during pre-spawn migration and spawning during the study period. In both study years, there was a sharp decline in mean egg numbers from spawning sites downstream of the rapids (439.7 eggs/2 min tow ± 990.6 SD) to upstream sites (5.9 eggs/2 min tow ± 19.4 SD). Physical barriers like rapids may reduce spawning habitat connectivity and could limit walleye production in the Maumee River.

Ohio↗
USGS2021

Lake-wide annual status of Mysis diluviana population in Lake Michigan in 2015

Mysis diluviana is one of the most abundant zooplankton by biomass in the Laurentian Great Lakes of North America, a predator of other zooplankton and an important prey for fishes. Studies of long-term trends in Lake Michigan have shown 2005–2016 densities to be 50–80% lower than 1990s densities, but these observations have been based on annual monitoring that is either spatially or seasonally limited. We combined Lake Michigan Mysis data from three annual programs and the 2015 Cooperative Science and Monitoring Initiative to achieve broad spatial coverage during spring, summer, and fall of 2015 and broad depth coverage during spring 2016. Lake-wide, annual density and biomass were 82 (SE: 10) Mysis /m 2 and 200 (SE: 36) mg dry mass/m 2 . Density and biomass estimates were highest offshore, generally higher in the north basin, and seasonally highest in summer. Annual lake-wide averages for depths >30 m were better captured by seasonally-extensive annual programs than spatially-extensive annual programs, although spring sampling may bias annual values low. Mysis cohorts grew 0.026 mm/d (age-0) and 0.007 to 0.027 mm/d (age-1). Annual mortality was 81–98%. Reproduction was fall-spring and seasonal lake-wide estimates ranged from 0.6 to 19.1% females brooding, 13–20 embryos/brood, and 3–46 embryos/m 2 . Annual production (423 mg dry mass/m 2 /yr, SE: 31) was lower than all but one previous estimate from lakes Michigan, Huron, and Ontario. While Mysis tend to persist, low Mysis production may be a concern for prey fishes that feed on Mysis .

Lake Michigan↗
USGS2021

Before the first meal: The elusive pre-feeding juvenile stage of the sea lamprey

Although sea lamprey ( Petromyzon marinus ) in the Laurentian Great Lakes have been studied intensively for more than 70 years, many questions remain about their complex life cycle. One of the least understood portions is the post-metamorphic stage (hereafter pre-feeding juvenile, PFJ) that occurs prior to parasitic feeding. PFJ must emerge from stream sediments and migrate downstream into nearshore feeding areas. Key uncertainties include the internal and exogenous triggers that regulate the timing and duration of the migration, and the mechanisms the animal uses to navigate, avoid predators, and locate their first host. However, many of these factors may vary predictably among natal streams in response to stable geomorphological and hydraulic characteristics that regulate the timing of movements (e.g., flood phenology), energetic costs (e.g., stream length), and risk (e.g., predator density). An improved understanding of the PFJ stage presents two opportunities to improve the success of sea lamprey control: (1) identification of streams where natural mortality during the PFJ stage is high, allowing for the reallocation of larval control to streams more likely to produce successful parasites, and (2) removal or killing of PFJs in streams where natural mortality is low. Either approach represents an opportunity to limit parasitic damage to valuable fish stocks. Here, we review the state of knowledge of the PFJ stage and identify critical knowledge gaps that, if addressed, could facilitate sea lamprey assessment and control by exploiting the behavior of PFJ as they outmigrate from streams in search of their first meal.

Journal of Great Lakes Research↗
USGS2021

Progress towards integrating an understanding of chemical ecology into sea lamprey control

The sea lamprey, Petromyzon marinus , is a destructive invader in the Laurentian Great Lakes that relies on several complex chemical cues to complete their life cycle. The central roles of chemical cues in sea lamprey reproduction provide opportunities to leverage knowledge of sea lamprey chemical ecology when developing alternative or supplemental strategies for sea lamprey control. A solid foundation has been laid regarding sea lamprey chemical ecology, with recent advances in our understanding of the migratory pheromone, male sex pheromone, and alarm cues broadening our fundamental understanding of the diversity, complexity, and evolution of chemical cues used by sea lamprey. Additionally, research applying semiochemicals in differing management scenarios has provided useful insights into the challenges of incorporating chemical cues into the sea lamprey control program. Here, we synthesize new findings related to fundamental research of chemosensory cues along with knowledge learned from management-based tests and explore options for integrating an understanding of chemical ecology into sea lamprey control in light of new knowledge. We also highlight current unknowns and future research needs that should be addressed prior to implementation of sea lamprey chemical ecology into the sea lamprey control program.

Great Lakes↗
USGS2021

Implications of tagging effects for interpreting the performance of sea lamprey traps in a large river

Abundance estimates can be crucial for managing species of economic concern. The accuracy of these estimates can depend on the methods used to track animals and to estimate abundance from tracking data. We tested experimentally if disparate estimates of trapping efficiency calculated for sea lamprey ( Petromyzon marinus ) in the St. Marys River near Sault Ste. Marie, Canada could be explained by effects related to the invasiveness and handling involved in tagging or the tag size used in the marking procedures. Trapping is used to gauge adult abundance, trapping efficiency, and success of a binational sea lamprey control program in the Laurentian Great Lakes, North America. Our experiment compared nightly catches of sea lamprey marked with external fin clips, surgically-implanted passive integrated transponder tags (PIT-only), and surgically-implanted PIT and acoustic tags (PIT+acoustic). We found no evidence that the probability of being trapped was affected by the added invasiveness and handling of internal tagging. Nightly recaptures of PIT-only tagged sea lamprey, relative to fin-clipped sea lamprey, were not different from expectations based on the numbers of individuals released from each treatment group. Conversely, there was evidence of effects related to tag size. Nightly recaptures of PIT+acoustic tagged sea lamprey, relative to PIT-only tagged sea lamprey, were lower than expected based on numbers of individuals released from each treatment group. Effects related to tag size partially explain the disparate estimates in trapping efficiency observed for sea lamprey.

Ontario↗
USGS2021

A renewed philosophy about supplemental sea lamprey controls

Invasive sea lamprey ( Petromyzon marinus ) populations in the Laurentian Great Lakes have been reduced by up to 90% through the use of selective pesticides (lampricides) and physical sea lamprey barriers that block spawning migrations. Nevertheless, other control methods are needed to achieve integrated pest management objectives, delay biological resistance, and address societal pressure to reduce pesticide use and restore lotic connectivity through dam removals. Despite decades of research and scientific advances, new control tools that focus on controlling adult and juvenile life stages have been rare because tactics have not been cost-effective alternatives to lampricides and sea lamprey barriers. Here, we propose a renewed philosophy highlighting that new control methods need not be true alternatives to lampricides and sea lamprey barriers (i.e., have similar effectiveness), but instead can be useful as supplemental controls integrated with current methods, especially in places where current methods are less effective due to environmental or societal conditions. Current case studies pairing multiple supplemental controls together on two Lake Huron tributaries, the Black Mallard and Cheboygan Rivers, have shown promise in reducing sea lamprey reproductive success, the scope of lampricide treatments, and ultimately the number of juvenile sea lampreys produced. Additional case studies are planned and will be evaluated within a decade-long adaptive assessment plan.

Journal of Great Lakes Research↗
USGS2021

What can commercial fishery data in the Great Lakes reveal about juvenile sea lamprey (Petromyzon marinus) ecology and management?

The Laurentian Great Lakes of North America support a large and profitable freshwater fishery, but one continuously beset by parasitism from the invasive sea lamprey ( Petromyzon marinus ). Despite being the life stage that inflicts damage to the fishery, therefore necessitating a bi-national control program, our knowledge of juvenile sea lamprey ecology is poor and their response to control efforts are not assessed. Incidental capture of juvenile sea lamprey by commercial fishers is one means to collect data on this enigmatic life stage, and in Lake Huron such data have been collated since 1967. Here, we explore incidental captures of juvenile sea lamprey and their hosts from northern Lake Huron between 1987 and 2017 ( n = 33,246 observations) to address four objectives. Firstly, we document collection efforts by fishers to provide historical context to the dataset. Secondly, we pose and test a series of questions related to fishery encounter, host selection, growth, distribution, and sex ratio to highlight how these types of data can be informative regarding juvenile sea lamprey ecology. Results presented here could be used to develop biological hypotheses to be addressed in future work. Thirdly, we directly assessed whether juvenile sea lamprey capture data could be useful in corroborating trends observed in adult sea lamprey abundance and wounding, as well as in identifying abundance and wounding hotspots. Lastly, we summarize research and outreach efforts that have benefited from the capture of juvenile sea lamprey in recent years.

Great Lakes↗
USGS2021

Environmental factors influencing annual sucker (Catostomus sp.) migration into a Great Lakes tributary

Fish migration in rivers is a growing area of concern as mounting anthropogenic influences, particularly fragmentation from dams and barriers, constitute major threats to global river species diversity. Barriers can impede the movement of fishes between areas critical to the completion of their lifecycle, affecting both population and ecosystem viability. In response, fish passage solutions have been identified as a critical need to maintain fisheries viability in the Laurentian Great Lakes, and around the world. Pivotal to the success of these fish passage solutions is a more complete understanding of the movement phenology and environmental cues that instigate migration. We used a dual-frequency identification sonar (DIDSON) to evaluate environmental triggers of river entry during spring and summer for three size classes of migratory fishes in the Boardman River, a Lake Michigan tributary. Our results indicate that medium size fish (>30 cm and < 50 cm), primarily composed of white sucker Catostomus commersonii and longnose sucker Catostomus catostomus were 21% more likely to enter the river at sunset and 25% less likely at midnight in comparison to midday. Entry rates of medium fish increased 6% for every 1 °C increase in river temperature, 4% for every 1 m 3 /s increase in river discharge from the day prior, and were reduced by 1% for every 10 cm increase in lake level. Understanding these processes in the tributaries of the Great Lakes is important to inform the fish passage solutions currently being developed for the Boardman River, and to inform management regulations for Great Lakes migratory fishes.

Michigan↗
USGS2021

Trade-offs between suppression and eradication of sea lampreys from the Great Lake

Ecosystem managers confronted with newly invasive species may respond with a program of suppression or eradication. Suppression of an invasive species refers to management of a species such that its effect on other biota in the local ecosystem is acceptable. Eradication is the removal of all individuals of a species from a defined region. We examine the cost and benefit trade-offs between suppression and eradication of Laurentian Great Lakes sea lampreys ( Petromyzon marinus ) based on discussions at the 3rd Sea Lamprey International Symposium (held in 2019). Substantial effort has been expended annually since the 1960s to suppress sea lampreys in the Great Lakes basin. Choosing between suppression and eradication is a value judgement, ideally made jointly by scientists, decision-makers, stakeholders, and society. Successful large-scale eradications have been limited to a small number of cases for which the cost to human society justified and supported the long-term commitment necessary for success. The greatest challenge to successful eradication of sea lampreys from the Great Lakes may be a suitable social, political, legal, and institutional environment. Preparations could be made now for a transition in which public pushback on current control methods (pesticide applications and barriers to fish passage) leads to more extensive use of an alternative control method, such as genetic control.

Great Lakes and Saint Lawrence River areas↗
USGS2021

Quantifying Great Lakes sea lamprey populations using an index of adults

Effective control of aquatic invasive species requires knowledge of the population throughout the infested area. Lake-wide assessments of invasive sea lampreys ( Petromyzon marinus ) are used to assess their status in the Laurentian Great Lakes, informing fisheries managers and decision makers in the sea lamprey control program. Initially these assessments focused on an estimate of absolute abundance, but later switched to an estimate of relative abundance as an index. In this paper, we describe the recently developed index of sea lamprey abundance and the reasons for its use. Rather than trying to estimate spawning run sizes of all Great Lakes tributaries, the index instead estimates run sizes of a small subset of index streams. Streams chosen for the index had large spawning runs and a history of trapping operations that consistently yielded mark-recapture estimates. This change enabled the sea lamprey control program to abandon a previously used regression model that predicted run size on streams with no sea lamprey traps. Further research is needed to determine how strongly correlated the index is with actual patterns in the lake-wide population of adult sea lampreys.

Lake Erie, Lake Huron, Lake Michigan, Lake Ontario↗
USGS2021

Trophic transfer efficiency in the Lake Superior food web: Assessing the impacts of non-native species

Ecosystem-based management relies on understanding how perturbations influence ecosystem structure and function (e.g., invasive species, exploitation, abiotic changes). However, data on unimpacted systems are scarce; therefore, we often rely on impacted systems to make inferences about ‘natural states.’ Among the Laurentian Great Lakes, Lake Superior provides a unique case study to address non-native species impacts because the food web is dominated by native species. Additionally, Lake Superior is both vertically (benthic versus pelagic) and horizontally (nearshore versus offshore) structured by depth, providing an opportunity to compare the function of these sub-food webs. We developed an updated Lake Superior EcoPath model using data from the 2005/2006 lake-wide multi-agency surveys covering multiple trophic levels. We then compared trophic transfer efficiency (TTE) to previously published EcoPath models. Finally, we compared ecosystem function of the 2005/2006 ecosystem to that with non-native linkages removed and compared native versus non-native species-specific approximations of TTE and trophic flow. Lake Superior was relatively efficient (TTE = 0.14) compared to systems reported in a global review (average TTE = 0.09), and the microbial loop was highly efficient (TTE > 0.20). Non-native species represented a very small proportion (<0.01%) of total biomass and were generally more efficient and had higher trophic flow compared to native species. Our results provide valuable insight into the importance of the microbial loop and represent a baseline estimate of non-native species impacts on Lake Superior. Finally, this work is a starting point for further model development to predict future changes in the Lake Superior ecosystem.

Lake Superior↗
USGS2021

Response of fish assemblages to restoration of rapids habitat in a Great Lakes connecting channel

Rapids habitats are critical spawning and nursery grounds for multiple Laurentian Great Lakes fishes of ecological importance such as lake sturgeon, walleye, and salmonids. However, river modifications have destroyed important rapids habitat in connecting channels by modifying flow profiles and removing large quantities of cobble and gravel that are preferred spawning substrates of several fish species. The conversion of rapids habitat to slow moving waters has altered fish assemblages and decreased the spawning success of lithophilic species. The St. Marys River is a Great Lakes connecting channel in which the majority of rapids habitat has been lost. However, rapids habitat was restored at the Little Rapids in 2016 to recover important spawning habitat in this river. During the restoration, flow and substrate were recovered to rapids habitat. We sampled the fish community (pre- and post-restoration), focusing on age-0 fishes in order to characterize the response of the fish assemblage to the restoration, particularly for species of importance (e.g. lake whitefish, walleye, Atlantic salmon). Following restoration, we observed a 40% increase in age-0 fish catch per unit effort , increased presence of rare species, and a shift in assemblage structure of age-0 fishes (higher relative abundance of Salmonidae, Cottidae, and Gasterosteidae). We also observed a “transition” period in 2017, in which the assemblage was markedly different from the pre- and post-restoration assemblages and was dominated by Catostomidae. Responses from target species were mixed, with increased Atlantic salmon abundance, first documented presence of walleye and no presence of lake sturgeon or Coregoninae.

Michigan, Ontario↗
USGS2022

Contemporary spatial extent and environmental drivers of larval coregonine distributions across Lake Ontario

Coregonine fishes are important to Laurentian Great Lakes food webs and fisheries and are central to basin-wide conservation initiatives. In Lake Ontario, binational management objectives include conserving and restoring spawning stocks of cisco ( Coregonus artedi ) and lake whitefish ( C. clupeaformis ), but the spatial extent of contemporary coregonine spawning habitat and the environmental factors regulating early life success are not well characterized. In Spring 2018, we conducted a binational ichthyoplankton assessment to describe the spatial extent of coregonine spawning habitat across Lake Ontario. We then quantified the relative importance of a suite of biophysical variables hypothesized to influence coregonine early life success using generalized additive mixed models and multimodel inference. Between April 10 and May 14, we conducted 1,092 ichthyoplankton tows and captured 2,350+ coregonine larvae across 17 sampling areas, predominantly within embayments. Although 95% of catches were in the eastern basin, coregonine larvae were also found in historical south shore spawning areas. Most coregonine larvae were cisco; <6% were lake whitefish. Observed catches of both species across sampling areas were strongly and similarly associated with ice cover duration, but the importance of site-specific characteristics varied, such as distance to shore and site depth for cisco and lake whitefish, respectively. These results suggest that regional-scale climatic drivers and local environmental habitat characteristics interact to regulate early life stage success. Furthermore, strong regional and cross-species variation in larval distributions emphasize the importance of lake-wide assessments for monitoring both the current eastern basin populations and potential expansions into western Lake Ontario habitats.

Lake Ontario↗
USGS2021

Towards improving an Area of Concern: Main-channel habitat rehabilitation priorities for the Maumee River

The Maumee River watershed in the Laurentian Great Lakes Basin has been impacted by decades of pollution and habitat modification due to human settlement and development. As such, the lower 35 km of the Maumee River and several smaller adjacent watersheds comprising over 2000 km 2 were designated the Maumee Area of Concern (AOC) under the revised Great Lakes Water Quality Agreement in 1987. As part of pre-rehabilitation assessments in the Maumee AOC, we assessed fish and invertebrate communities in river km 24–11 of the Maumee River to identify: 1) areas that exhibit the highest biodiversity, 2) habitat characteristics associated with high biodiversity areas, 3) areas in need of protection from further degradation, and 4) areas that could feasibly be rehabilitated to increase biodiversity. Based on benthic trawl data, shallow water habitats surrounding large island complexes had the highest fish diversity and catch per unit effort (CPUE). Electrofishing displayed similar fish diversity and CPUE patterns across habitat types early in the study but yielded no discernable fish diversity or CPUE patterns towards the end of our study. Although highly variable among study sites, macroinvertebrate density was greatest in shallow water habitats <2.5 m and around large island complexes. Our results provide valuable baseline data that could act as a foundation for developing rehabilitation strategies in the lower Maumee River and for assessing the effectiveness of future aquatic habitat rehabilitation projects. In addition to increasing in-channel habitat, watershed-scale improvements of water quality might be necessary to ensure rehabilitation strategies are successful.

Ohio↗
USGS2021

Distribution, abundance and spatial variability of microplastic pollution on the surface of Lake Superior

In 2014, 94 paired neuston net samples (0.5 mm mesh) were collected from the surface waters of Lake Superior. These samples comprise the most comprehensive surface water survey for microplastics of any of the Great Lakes to date, and the first to employ double net trawls. Microplastic abundance estimates showed wide variability, ranging between 4000 to more than 100,000 particles/km 2 with most locations having abundances between 20,000 to 50,000 particles/km 2 . The average abundance in Lake Superior was ~30,000 particles/km 2 which was similar to previous estimates within this Laurentian Great Lake and suggests a total count of more than 2.4 billion (1.7 to 3.3 billion, 95% confidence interval) particles across the lake’s surface. Distributions of plastic particles, characterized by size fraction and type, differed between nearshore and offshore samples, and between samples collected in the eastern versus western portion of the lake. Most of the particles found were fibers (67%), and most (62%) were contained in the smallest classified size fraction (0.50–1 mm). The most common type of polymer found was polyethylene (51%), followed by polypropylene (19%). This is consistent with global plastics production and results obtained from other studies. No statistically significant difference was detected between the paired net samples, indicating that single net sampling should produce a representative estimate of microplastic particle abundance and distribution within a body of water.

Lake Superior↗