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Brian Weidel

Publications and source records attributed to Brian Weidel.

At least 55 records · Page 3Linked to original sources
USGS2021

Incorporation of non-native species in the diets of cisco (Coregonus artedi) from eastern Lake Ontario

Cisco Coregonus artedi was once an important native fish in Lake Ontario; however, after multiple population crashes, the cisco stock has yet to recover to historic abundances. Rehabilitation of cisco in Lake Ontario is a fish community management objective, but the extent to which recent non-native species and pelagic food web changes have influenced cisco is not well understood. We described cisco diets in contemporary Lake Ontario following the addition and spread of non-native zooplankton species. We collected 618 cisco and processed 178 for full diet analysis in eastern Lake Ontario using mid-water trawls and bottom-set gill nets from 2016 to 2020. We found that Lake Ontario cisco were mostly zooplanktivorous, and non-native zooplankton dominated their diet during July and September. Cisco smaller than 300 mm had a more diverse diet including both native and non-native zooplankton, while cisco larger than 300 mm fed almost exclusively on non-native predatory cladocerans Bythotrephes longimanus and Cercopagis pengoi (98.9% consumed prey dry mass). We also found fish eggs, presumed to be of coregonine origin in 75% of non-empty December-collected cisco diets, suggesting eggs subsidize cisco diets when available. Juvenile round goby Neogobius melanostomus , alewife Alosa pseudoharengus and rainbow smelt Osmerus mordax were found in 2% of all analyzed non-empty stomachs. Lake Ontario cisco diet appears to be more similar to zooplanktivorous Lake Superior cisco than Lake Michigan where piscivory is prevalent. Lake Ontario cisco diets reflected zooplankton community changes indicating that non-native predatory cladocerans are now an important energy source supporting this native species.

Eastern Lake Ontario↗
USGS2021

Slimy sculpin depth shifts and habitat squeeze following the round goby invasion in the Laurentian Great Lakes

The collapse of Diporeia spp. and invasions of dreissenid mussels (zebra, Dreissena polymorpha ; quagga, D. bugensis ) and round goby ( Neogobius melanostomus ) have been associated with declines in abundance of native benthic fishes in the Great Lakes, including historically abundant slimy sculpin ( Cottus cognatus ). We hypothesized that as round goby colonized deeper habitat, slimy sculpin avoided habitat competition, predation, and aggression from round goby by shifting to deeper habitat. Accordingly, we predicted increased depth overlap of slimy sculpin with both round goby and deepwater sculpin ( Myoxocephalus thompsonii ) that resulted in habitat squeeze by both species. We used long-term bottom trawl data from Lakes Michigan, Huron, and Ontario to evaluate shifts in slimy sculpin depth and their depth overlap with round goby and deepwater sculpin. Lake Huron most supported our hypotheses as slimy sculpin shifted to deeper habitat coincident with the round goby invasion, and depth overlap between slimy sculpin and both species recently increased. Slimy sculpin depth trends in Lakes Michigan and Ontario suggest other ecological and environmental factors better predicted sculpin depth in these lakes.

Lake Huron, Lake Michigan, Lake Ontario↗
USGS2021

Lake Ontario cisco population dynamics based on long-term surveys

Prior to European settlement, cisco (Coregonus artedi) were likely one of Lake Ontario’s most abundant fishes but currently represent a small portion of the fish community. To understand how the population has changed over the past 70 years we compared trends in annual catch rates from gillnet and bottom trawl surveys and commercial fishery landings. In surveys, cisco were generally rare, and represented 0.2, 0.4, and 0.001% of all fish caught in two gillnet surveys and bottom trawl surveys. Cisco catch rates in gillnets and trawls were positively correlated and correlations increased when gillnet catches two years later were compared to trawls since trawls tended to capture smaller, juvenile-sized cisco relative to gillnets. Survey catch rates suggest recruitment is generally low, but discrete periods of relatively greater recruitment in the 1980s and mid-2010s suggest reproductive conditions for cisco vary temporally. Trawl surveys were the most spatially extensive survey and illustrated catch rates were highest in northeastern Lake Ontario. Greater cisco abundance in this region may be related to more-abundant embayment spawning habitat, greater distance from winter aggregations of nonnative planktivores, or more appropriate environmental conditions during spawing. At the basin scale, Lake Ontario bottom trawl catch per unit effort (CPUE) was positively correlated to Lake Superior trawl CPUE suggesting a regional driver, such as climate, may be similarly impacting both populations. Concurrent patterns across Lake Ontario surveys support the idea that cisco are currently a small portion of the fish community, recruitment remains inconsistent, and habitats in northeastern Lake Ontario appear critical to the remnant populations.

Lake Ontario↗
USGS2021

Lake trout rehabilitation in Lake Ontario, 2020

Each year we report on the progress toward rehabilitation of the Lake Ontario lake trout (Salvelinus namaycush) population, including the results of stocking, annual assessment surveys, creel surveys, and evidence of natural reproduction observed from standard surveys performed by USGS and NYSDEC. Response to the COVID-19 pandemic limited survey effort such that spring and summer bottom trawl surveys and the creel survey were not completed in 2020, and sites sampled during the fall gillnet survey were limited to those east of Rochester, NY. The catch per unit effort of adult lake trout in gill nets increased each year from 2008-2014, recovering from historic lows recorded during 2005-2007. Adult abundances declined each year from 2015 to 2017; and in 2017 were about 35% below the 2014 peak and 17% below the 1999-2004 mean. Adult abundance increased in 2018 by 51% over the 2017 value and remained nearly stable between 2018 and 2020. The 2020 rate of wounding by sea lamprey (Petromyzon marinus) on lake trout caught in gill nets was 2.27 A1 wounds (fresh wound) per 100 lake trout and was near target (2 wounds per 100 lake trout). Condition values for adult lake trout, indexed in September from the predicted weight for a 700 mm lake trout from annual length-weight regressions and Fulton’s K for age-6 males, were among the highest levels observed for the 1983-2020 time series. Reproductive potential for the adult stock indexed from the CPUE of mature females ≥ 4000 g was again above the target in 2020 continuing a trend observed in nine of the last ten years. The 2020 catch of young wild lake trout marked the 26th observation in the last 27 years, however the low numbers of native adults observed during that time period continues to indicate substantial restoration impediments still exist.

Lake Ontario↗
USGS2021

2020 Status of the Lake Ontario lower trophic levels

Significant Findings for Year 2020: Note that due to covid-19 restrictions, offshore sampling was limited in 2020. 1) May – Oct total phosphorus (TP) in 2020 was 10.6 µg/L (offshore) and 7.7 µg/L (nearshore), higher than the long-term (1995-2019) average in the offshore (6.2 µg/L) and close to average in the nearshore (7.8 µg/L); mean TP values for the past decade (2010-2019) were 6.0 µg/L and 7.9 µg/L in the offshore and nearshore, respectively. In 2020, TP concentrations were significantly higher (p=0.03) in the offshore compared to the nearshore. Note that offshore duplicate samples had high relative percent difference (average 54%. 6-117%) making inferences for the offshore in 2020 uncertain. 2) May – Oct epilimnetic chlorophyll-a was similar at nearshore (1.9 µg/L) and offshore (2.1 µg/L) sites. These values were slightly higher than the average for 1995 – 2019 (1.7 µg/L, offshore; 1.5 µg/L, nearshore) and higher than for the last decade (1.4 µg/L, both offshore and nearshore). 3) May – Oct Secchi depth ranged from 3.5 m to 10.4 m (11 ft to 34 ft) at individual sites and was not significantly different between nearshore (6.3 m; 20.7 ft) and offshore (6.5 m; 21.3 ft) locations. Long-term (1995-2019) average was 7.2 m in the offshore and 6.4 m in the nearshore; means for the last decade were 7.8 m in the offshore and 6.2 m in the nearshore. 4) Despite higher TP values in 2020 than in recent years, TP, chlorophyll-a and Secchi depth are indicative of oligotrophic conditions in the offshore of Lake Ontario. 5) Nearshore summer zooplankton biomass was 10.5 µg/L, near the all-time low (9.4 µg/L, 2017) since monitoring began in 1995. Offshore epilimnetic summer zooplankton biomass was 11.7 µg/L. These values are similar to biomass in the last decade (2010-2019). 6) Peak (July) epilimnetic biomass of Cercopagis was 3.0 µg/L in the nearshore and represented 25% of the zooplankton community at that time; Cercopagis was absent from the July offshore epilimnetic samples in 2020 but was present in whole water column samples taken in August by other agencies. Epilimnetic biomass of Bythotrephes peaked in late-September in both the nearshore (1.0 µg/L) and offshore (1.8 µg/L) and represented 10% and 18% of the zooplankton community at those times, respectively. 7) Summer nearshore and offshore epilimnetic zooplankton density and biomass declined significantly 1995 – 2020. The declines were due mainly to reductions in cyclopoid copepods in both habitats.

Lake Ontario↗
USGS2021

Lake Ontario April prey fish survey and Alewife assessment, 2021

The Lake Ontario April bottom trawl survey and Alewife, Alosa psuedoharengus population assessment are conducted annually to track prey fish community status and aid management decisions related to predator-prey balance. No survey was conducted in 2020 due to the Covid-19 pandemic. The 2021 survey included 248 bottom trawls in both U.S. and Canadian waters, from March 30 - May 7 in the main lake and embayment regions, at depths ranging from 5 – 221 m (16 - 729 ft). The survey captured 947,102 fish, from 30 species with a total weight of 9,191 kg (20,220 lbs). Alewife were 89.2% of the catch by number while Rainbow Smelt, Osmerus mordax , Round Goby Neogobius melanostomus , and Deepwater Sculpin Myoxocephalus thompsonii comprised 5.6, 2.3, and 1.7% of the catch, respectively. Rainbow Smelt biomass in 2021 was among the highest values observed since 1997, especially in U.S. waters. The biomass index for Cisco, Coregonus artedii also increased, primarily due to catches and greater survey effort in the Bay of Quinte. Threespine stickleback, Gasterosteus aculeatus and Emerald Shiner, Notropis atherinoides biomasses remain low. No Bloater, Coregonus hoyi were captured during the 2021 survey. In 2021, the lake-wide Alewife biomass index increased substantially from 2019 due to the presence of an exceptionally high catch of age-1 Alewife (2020 year class). The biomass index of adult Alewife (age-2 and up) declined slightly since 2019, which was expected since Alewife reproduction was generally below average from 2016 to 2019. Expanding the survey spatial extent from U.S. waters to a lake-wide survey in 2016 has improved our ability to estimate Alewife survival and has provided more accurate estimates of Lake Ontario Alewife biomass and density. Simulation modeling based on recent estimates of survival, growth, and reproduction suggests the adult Alewife biomass will likely increase in 2022 and 2023. As part of a continued effort to improve prey fish surveys, we employed hydroacoustic sampling during the 2021 April trawl survey to estimate fish densities in open-water, pelagic habitats not sampled by the bottom trawl. We found fish density, in waters above the trawl headline depth (3m off bottom to surface), were approximately ~100 times lower than pelagic prey fish densities from bottom trawls. These results support the idea that at this time of year, when the warmest water is on the lake bottom, Alewife and most other prey fish primarily inhabit deep, near bottom regions and can be effectively sampled with bottom trawls. We were not able to apportion acoustics targets to species, however the low mean target strength (-43 decibels, dB) suggested these were small fishes (e.g., 100 mm). The greatest hydroacoustics densities were found near the Niagara River confluence and future surveys may use midwater trawls to determine which species these were and continue to improve this multi-agency survey.

Lake Ontario↗
USGS2021

Angler-caught salmonid diets illustrate Lake Ontario Alewife population and predator-prey dynamics

Lake Ontario fisheries decision makers use information about the status and trajectory of prey fish populations, such as alewife, for salmonid sport fish management. In 2020, the April bottom trawl survey, typically used to assess alewife, was canceled the day after it began due to Coronavirus health concerns. This prompted NYSDEC and USGS science teams to initiate a diet study using angler-collected samples to provide information on the alewife population in 2020. Over 800 salmonid diets were collected from across Lake Ontario southern shore from May through September. Past diet studies illustrated alewife size distribution in piscivore diets overlapped April trawl survey-based distributions (proportion overlap = 0.51- 0.81, n = 5). In 2020 few age-1 sized alewife (40 - 140 mm) were observed in salmonid diets, suggesting that 2019 alewife reproduction was below average. Given that alewife reproductive success from 2017 and 2018 were also below average, the 2021 Lake Ontario adult alewife population will likely be at a record low for the 43-year time series. For all years and piscivore species, adult-sized Alewife ( total length > 140 mm) were the predominate size consumed by piscivores greater than 400 mm (~ 16 inches). Sizes of alewife in the diet were positively related to piscivore size. Diet composition, by dry weight, indicated alewife remain the dominant prey supporting salmonids and diets were generally similar to trawl-based estimates of prey fish community composition. The most notable deviation from this pattern included round goby, which comprised substantial portions of brown trout and lake trout diets in 2013 and 2020. These results inform decision making related to Lake Ontario salmonid stocking and predator prey dynamics and highlight the utility of angler-caught piscivore diets for indicating alewife population status.

Lake Ontario↗
USGS2020

Comparison of specimen- and image-based morphometrics in Cisco

Morphometric data from fish are typically generated using one of two methods: direct measurements made on a specimen or extraction of distances from a digital picture. We compared data on 12 morphometrics collected with these two methods on the same collection of Cisco Coregonus artedi from Lake Ontario, North America, to assess the degree of bias in measurements made directly on a specimen- vs. an image-based method. We also assessed the degree of reproducibility within the image-based method by evaluating the amount of variation between different analysts for each morphometric method. Our results indicate specific morphometrics may be more prone to bias across the two methods and between analysts. Four of 12 morphometrics evaluated showed significant deviation from a 1:1 relationship that would be expected if the imaged-based method produced accurate specimen-based measurements. Pelvic fin length and pelvic–anal fin distance had the highest between-analyst variation for image-based landmarks, indicating low reproducibility for these metrics, compared with pectoral fin or total length, which had lower between-analyst variation. Although some morphometric measurements can be accurately obtained with either method, and therefore potentially used interchangeably in studies on Cisco morphology, our findings highlight the importance of considering method bias in morphometric studies that use data collected by different methods.

Journal of Fish and Wildlife Management↗
USGS2020

A century of intermittent eco‐evolutionary feedbacks resulted in novel trait combinations in invasive Great Lakes alewives (Alosa pseudoharengus)

Species introductions provide opportunities to quantify rates and patterns of evolutionary change in response to novel environments. Alewives ( Alosa pseudoharengus ) are native to the East Coast of North America where they ascend coastal rivers to spawn in lakes and then return to the ocean. Some populations have become landlocked within the last 350 years and diverged phenotypically from their ancestral marine population. More recently, alewives were introduced to the Laurentian Great Lakes (~150 years ago), but these populations have not been compared to East Coast anadromous and landlocked populations. We quantified 95 years of evolution in foraging traits and overall body shape of Great Lakes alewives and compared patterns of phenotypic evolution of Great Lakes alewives to East Coast anadromous and landlocked populations. Our results suggest that gill raker spacing in Great Lakes alewives has evolved in a dynamic pattern that is consistent with responses to strong but intermittent eco‐evolutionary feedbacks with zooplankton size. Following their initial colonization of Lakes Ontario and Michigan, dense alewife populations likely depleted large‐bodied zooplankton, which drove a decrease in alewife gill raker spacing. However, the introduction of large, non‐native zooplankton to the Great Lakes in later decades resulted in an increase in gill raker spacing, and present‐day Great Lakes alewives have gill raker spacing patterns that are similar to the ancestral East Coast anadromous population. Conversely, contemporary Great Lakes alewife populations possess a gape width consistent with East Coast landlocked populations. Body shape showed remarkable parallel evolution with East Coast landlocked populations, likely due to a shared response to the loss of long‐distance movement or migrations. Our results suggest the colonization of a new environment and cessation of migration can result in rapid parallel evolution in some traits, but contingency also plays a role, and a dynamic ecosystem can also yield novel trait combinations.

Great Lakes↗
USGS2020

Benthic habitat is an integral part of freshwater Mysis ecology

Diel vertical migration (DVM) is common in aquatic organisms. The trade‐off between reduced predation risk in deeper, darker waters during the day and increased foraging opportunities closer to the surface at night is a leading hypothesis for DVM behaviour. Diel vertical migration behaviour has dominated research and assessment frameworks for Mysis , an omnivorous mid‐trophic level macroinvertebrate that exhibits strong DVM between benthic and pelagic habitats and plays key roles in many deep lake ecosystems. However, some historical literature and more recent evidence indicate that mysids also remain on the bottom at night, counter to expectations of DVM. We surveyed the freshwater Mysis literature using Web of Science (WoS; 1945–2019) to quantify the frequency of studies on demographics, diets, and feeding experiments that considered, assessed, or included Mysis that did not migrate vertically but remained in benthic habitats. We supplemented our WoS survey with literature searches for relevant papers published prior to 1945, journal articles and theses not listed in WoS, and additional references known to the authors but missing from WoS (e.g. only 47% of the papers used to evaluate in situ diets were identified by WoS). Results from the survey suggest that relatively little attention has been paid to the benthic components of Mysis ecology. Moreover, the literature suggests that reliance on Mysis sampling protocols using pelagic gear at night provides an incomplete picture of Mysis populations and their role in ecosystem structure and function. We summarise current knowledge of Mysis DVM and provide an expanded framework that more fully considers the role of benthic habitat. Acknowledging benthic habitat as an integral part of Mysis ecology will enable research to better understand the role of Mysis in food web processes.

Freshwater Biology↗
USGS2020

A synthesis of the biology and ecology of sculpin species in the Laurentian Great Lakes and implications for the adaptive capacity of the benthic ecosystem

The Laurentian Great Lakes have experienced recent ecosystem changes that could lead to reductions in adaptive capacity and ultimately a loss of biodiversity and production throughout the food web. Observed changes in Great Lakes benthic communities include declines of native species and widespread success of invasive species like dreissenid mussels in all but Lake Superior. Understanding the ecology of native benthic deepwater preyfish and the reasons for their declines is important for predicting future losses in adaptive capacity and diversity, as well as managing the Great Lakes ecosystem to avoid such losses. Native sculpin species ( Cottus bairdii , C. cognatus , C. ricei , Myoxocephalus thompsonii ) historically were among the most abundant of the Great Lakes native deepwater benthic preyfish community and are an important link between offshore benthic and pelagic food webs. With one exception, these species have declined in abundance throughout the Great Lakes in recent years, but relatively little is known about their biology and ecology. This review synthesizes the available knowledge for the Great Lakes sculpin species and provides suggestions for future research efforts, which include understanding reproductive ecology and spawning behavior, connectivity and dispersal of populations, early life history, and influences of interactions with native and non-native species.

Great Lakes↗
USGS2020

Differentiation between lake whitefish and cisco eggs based on diameter

Cisco ( Coregonus artedi ) and lake whitefish ( Coregonus clupeaformis ) are native fish species of management concern in the Laurentian Great Lakes that often overlap in spawning locations and timing. Thus, species-level inference from in situ sampling requires methods to differentiate their eggs. Genetic barcoding and hatching eggs to visually identify larvae are used but can be time and cost intensive. Observations in published literature indicate that lake whitefish eggs may be larger than cisco eggs in the Great Lakes, but this has not yet been substantiated. Samples from shared spawning grounds are unlikely to contain similarly sized or colored eggs from other species. Thus, we assessed whether lake whitefish and cisco eggs could be separated based on size alone. Fertilized, hardened eggs were collected in situ during spawning at Elk Rapids, Lake Michigan and Chaumont Bay, Lake Ontario and preserved in ethanol. Individual eggs were measured and genetically identified. Mean diameter for cisco (2.45 mm, SD = 0.22, n = 444) was smaller than for lake whitefish (3.21 mm, SD = 0.20, n = 99). We used classification trees to identify a species-separating size threshold of 2.88 mm (95% bootstrap CI = [2.877, 2.976]), which classified eggs with an accuracy rate of 96%. Differences between species across other samples from the same locations were mostly consistent with the threshold size, but we suggest validation if applying this method to other populations. Separation of cisco and lake whitefish eggs by diameter can be accurate, efficient, and especially suitable for large sample sizes.

Lake Michigan, Lake Ontario↗
USGS2020

Lake Ontario deepwater sculpin recovery: An unexpected outcome of ecosystem change

Fish population recoveries can result from ecosystem change in the absence of targeted restoration actions. In Lake Ontario, native Deepwater Sculpin Myoxecephalus thompsonii, were common in the late-1800s, but by the mid-1900s the species was possibly extirpated. During this period mineral nutrient inputs increased and piscivore abundance declined, which increased the abundance of the nonnative planktivores Alewife Alosa pseudoharengus and Rainbow Smelt Osmerus mordax. Deepwater Sculpin larvae are pelagic and vulnerable to predation by planktivores. Annual bottom trawl surveys did not capture Deepwater Sculpin from 1978 – 1995 (n = 6,666 tows) despite sampling appropriate habitat (trawl depths: 7 – 170 m). The absence of observations during this time resulted in an elevated conservation status for the species, but no restoration actions were initiated. In 1996, three individuals were caught in bottom trawls, the first observed since 1972. Since then, their abundance has increased and in 2017 they were the second most abundant Lake Ontario prey fish. The food-web changes that occurred from 1970 through the 1990s contributed to this recovery. Alewife and Rainbow Smelt abundance declined during this period due to predation by stocked salmonids and legislation that reduced nutrient inputs and food web productivity. In the 1990s, the proliferation of nonnative, filter-feeding dreissenid mussels dramatically increased water clarity. As light penetration increased, the early-spring depth distribution of Alewife and Rainbow Smelt shifted deeper, away from larval Deepwater Sculpin habitat. The intentional and unintentional changes that occurred in Lake Ontario were not targeted at Deepwater Sculpin restoration but resulted in conditions that favored the species’ recovery. While standard surveys documented the recovery, more diverse information (e.g. observations in deep habitats and early-life stages) would have improved our understanding of why the species recovered when it did. Annual Lake Ontario trawl surveys have collaboratively expanded their spatial extent and diversified habitat sampled, based on lessons learned from the Deepwater Sculpin recovery.

Lake Ontario↗
USGS2020

Bottom trawl assessment of Lake Ontario prey fishes, 2019

Multi-agency, collaborative Lake Ontario bottom trawl surveys provide information for decision making related to Fish Community Objectives including predator-prey balance and understanding prey fish community diversity. In 2019, bottom trawl surveys in April (n = 252 tows) and October (n = 160 tows) sampled main lake and embayments at depths from 5–226 m. Combined, the surveys captured 283,383 fish from 39 species. Alewife were 67% of the total catch by number while round goby, deepwater sculpin, and rainbow smelt comprised 13, 10, and 4% of the catch, respectively. In 2019, the lake-wide adult alewife biomass index declined from 2018 and age-1 biomass, a measure of reproductive success the previous year, was low. Year-class catch curve models identified years where estimates from surveys conducted only in U.S. waters were biased, potentially due to a greater portion of the alewife population inhabiting unsampled Canadian waters. Accounting for spatial survey bias, these model estimates indicated the 2019 adult alewife biomass was the lowest value in the 42-year time series. Models also identified the extent to which age-1 alewife biomass was historically underestimated, however lake-wide results from 2016-2019 appear less biased. If below-average year-class estimates from 2017 and 2018 are accurate, adult alewife biomass will continue to decline in 2020. Abundance indices for other pelagic prey fishes such as rainbow smelt, threespine stickleback, emerald shiner, and cisco were low and similar to 2018 values. Pelagic prey fish diversity is low because a single species, alewife, dominates the community. Deepwater sculpin and round goby were the most abundant demersal (bottom-oriented) prey fishes in 2019. Despite declines in slimy sculpin and other nearshore prey fishes, demersal prey fish community diversity has increased as deepwater sculpin and round goby comprise more even portions of the community. New experimental trawl sites in embayment habitats generally captured more species, a higher proportion of native species, and higher densities relative to main lake habitats. In 2019, a western tubenose goby (Proterorhinus semilunaris) was captured for the first time in the trawl surveys.

Lake Ontario↗
USGS2020

2019 Status of the Lake Ontario Lower Trophic Levels

Spring total phosphorus (TP) in 2019 was 3.2 µg/L (offshore) and 4.7 µg/L (nearshore), both all-time lows; however, there is no significant time trend in our data series (1995-2019 for nearshore; 2002-2019 for offshore). Apr/May – Oct mean TP concentrations were low at both nearshore and offshore locations (range, 3.7 – 6.5 µg/L). TP and SRP concentrations were not significantly different between nearshore and offshore habitats. Chlorophyll- a and Secchi depth values are indicative of oligotrophic conditions in nearshore and offshore habitats. Offshore summer chlorophyll-a declined significantly 1995 – 2019. Nearshore chlorophyll- a increased 1995 – 2004 and then stabilized 2005 – 2019. In 2019, epilimnetic chlorophyll-a averaged between 1.3 and 2.9 μg/L across sites, and Apr/May – Oct concentrations were not significantly different between nearshore and offshore sites. Summer Secchi depth increased significantly in the offshore 1995 – 2019 from ~6 m to ~8 m. In the nearshore Secchi depth increased 1995 – 2004 but has remained around 6 m since 1999. Apr/May – Oct Secchi depth ranged from 3.8 m to 9.1 m (12 ft to 30 ft) at individual sites and was significantly higher offshore (7.6 m; 25 ft) than nearshore (5.7 m; 19 ft). In 2019, nearshore summer zooplankton biomass increased to 16.7 mg/m 3 after an all-time low (10.3 mg/m 3 ) in 2017. Offshore biomass (12.0 mg/m 3 ) was near the all-time low (8.1 mg/m 3 , 2006). Apr/May – Oct epilimnetic zooplankton density and biomass were not different between nearshore and offshore sites. However, zooplankton average size was significantly higher in the offshore (0.72 mm) than the nearshore (0.61 mm). Peak (July) epilimnetic biomass of Cercopagis was 2.4 mg/m 3 in the nearshore and 1.4 mg/m 3 in the offshore. Peak (September) epilimnetic biomass of Bythotrephes was 2.0 mg/m 3 in the nearshore and 2.9 mg/m 3 in the offshore. Summer nearshore zooplankton density and biomass declined significantly 1995 – 2004 and then remained stable 2005 – 2019. The decline was due mainly to reductions in cyclopoids copepods. Summer epilimnetic daytime offshore zooplankton density decreased significantly 1995 – 2004, but biomass did not. Density and biomass declined significantly 1995 – 2019. Density was 3885/m 3 in 2019, about one-fourth the level observed the previous year. Offshore summer epilimnetic zooplankton biomass in 2019 was 12 mg/m 3 —well below the mean from 2005 – 2018 (20 mg/m 3 ). Most offshore zooplankton biomass was found in the metalimnion in July and early-October, and in the hypolimnion in September. Limnocalanus dominated the metalimnion in July while daphnids comprised most of the biomass in October. In September, Limnocalanus dominated the hypolimnion.

New York, Ontario↗