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Jeremy P. Holden

Publications and source records attributed to Jeremy P. Holden.

At least 19 recordsLinked to original sources
USGS2026

Evaluating the use of uncrewed surface vessels to enhance Lake Erie acoustic prey-fish surveys

Incorporation of autonomous uncrewed surface vessels (USVs) into large-scale acoustic surveys may enhance spatiotemporal extent and quality of fish density estimates. Lake Erie is currently surveyed by three motorized research vessels (RVs), which annually collect acoustic data and estimate prey-fish abundances. To evaluate the feasibility of incorporating a USV into the existing survey to increase spatial coverage, we compared paired acoustic measurements of target strength (TS), area backscattering coefficient (ABC), and areal fish density from four motorized RVs (three currently and one historically used) and one USV (Saildrone Explorer). Acoustic data were collected along 2-km transects during five sampling events distributed throughout Lake Erie. Despite observing event-specific differences that were likely impacted by sampling conditions, survey-level estimates of areal density between RVs and the USV were comparable (i.e., RV 6% lower than USV). RVs and the USV appear to react differently (speed and stability) to variable environmental conditions (wind and waves), which impacted data quality and must be mitigated in future applications. However, similar to previous comparisons in marine and freshwater environments, incorporating a USV into an acoustic survey could enhance annual fish density estimates by increasing effort, spatial coverage, logistical flexibility, and biological data collection opportunities from RVs.

Michigan, New York, Ohio, Pennsylvania↗
USGS2025

Lake Ontario spring prey fish bottom trawl survey and Alewife assessment, 2025

The multi-agency Lake Ontario spring prey fish survey quantifies changes in pelagic prey fish populations, in particular Alewife Alosa pseudoharengus , which are the primary prey supporting the lake’s sport fishes. The 2025 survey included 230 trawls in the main lake and embayments and sampled depths from 5.5 to 245 m (15 – 810 ft). The survey captured 504,541 fish from 33 species with a total weight of 7,301 kg (16,095 lbs). Alewife were 85% of the total catch numerically, while Yellow Perch Perca flavescens , Round Goby Neogobius melanostomus , Deepwater Sculpin Myoxocephalus thompsonii , and Rainbow Smelt Osmerus mordax , comprised 5%, 4%, 3%, and 1% of the catch, respectively. The Alewife biomass index decreased from 2024 to 2025 (83 to 78 kg·ha -1 ) however due to an abundant 2024 Alewife year class the density index increased from 3,727 to 9,182 fish per ha -1 . The Age-1 biomass (2024 year class) was 27.5 kg·ha -1 , which was the greatest value estimated in the modern time series (since 1997). The abundance estimate for the 2024 Alewife year class (13.8 billion) was more than three times the number of all other Alewife combined (3.6 billion). Adult Alewife abundance decreased in 2025 which was consistent with predictions from 2024. Those predictive models suggested that adult Alewife biomass is likely to increase in 2026 and 2027, as the 2024 year class matures. Alewife condition declined in 2025, which was expected given the relatively high Alewife density. Acoustic-based prey fish densities were greater than previous years acoustic estimates especially at depths from 180 – 220 m (591 – 722 ft), however acoustic based densities continue to be substantially lower than trawl-based densities. The 2025 biomass index was similar to 2024 for Emerald Shiner Notropis atherinoides and Threespine Stickleback Gasterosteus aculeatus , but was lower for Rainbow Smelt, and higher for Cisco Coregonus artedi . Three purported Bloater Coregonus hoyi were caught in the 2025 survey. Analysis of archived tissue identified five Bloater captured in previous surveys which increased the total number caught in Lake Ontario bottom trawl surveys to n = 24, since restoration stocking began in 2012. Whole lake density estimates of Lake Whitefish Coregonus clupeaformis increased in 2025 relative to 2024. Those density increases were due to increased catches in Canadian waters, as density in U.S. waters has remained low. The density index for wild or naturally reproduced juvenile Lake Trout Salvelinus namaycush increased in 2025 relative to 2024, with the most frequent catches occurring in waters around the Niagara River.

Lake Ontario↗
USGS2024

Bottom trawl assessment of Lake Ontario's benthic preyfish community, 2023

Since 1978, surveys of Lake Ontario preyfish communities have provided information on the status and trends of the benthic preyfish community related to Fish Community Objectives that includes understanding preyfish population dynamics and community diversity. Beginning in 2015, the benthic preyfish survey expanded from US-only to incorporate Canadian sites, increasing the survey’s spatial coverage to a lake-wide scale. Additionally, sampling in eastern US embayments (Black River, Chaumont, Guffin, and Henderson Bays), that were historically sampled during a September bottom trawl survey to index Yellow Perch (Perca flavescens; 1978–2007), resumed in 2015. The current survey provides abundance indices for sculpins, Round Goby (Neogobius melanostomus) and Bloater (Coregonus hoyi) with survey techniques, gear and timing comparable to Lake Michigan. This alignment provides a necessary biological reference point for measuring the success of Lake Ontario Bloater reintroduction. In 2023, the collaborative benthic preyfish survey completed 188 bottom trawl tows across main lake and embayment sites at depths from 6 to 249 m. In total, the 2023 survey sampled 85,801 fish from 38 species. Round Goby was the most common species comprising 43% of the total catch, followed by Deepwater Sculpin (Myoxocephalus thompsonii), and Alewife (Alosa pseudoharengus) at 22% and 13%, respectively. Slimy Sculpin (Cottus cognatus) lake-wide biomass density (0.06 kg/ha) remained low relative to historical observations from US waters during the 1980-1990s and was orders of magnitude lower in US than Canadian waters. Lake-wide Deepwater Sculpin biomass density remains high since the population recovery (4.1 kg/ha). Embayment catches continue to have unique species assemblages compared to main lake habitat. Historically common native benthic preyfish species like Trout-perch (Percopsis omiscomaycus), Spottail Shiner (Notropis hudsonius), and darters (Etheostoma spp.), that are now rare at main lake trawl sites, still occur in some embayment trawl sites.

Lake Ontario↗
USGS2024

Lake Ontario April prey fish survey results and Alewife assessment, 2024

The Lake Ontario April bottom trawl survey assesses pelagic prey fish populations, in particular Alewife Alosa pseudoharengus , which are the primary prey supporting the lake’s sport fish populations. The 2024 survey included 234 trawls in the main lake and embayments and sampled depths from 3.9 to 245 m (13 – 809 ft). The survey captured 441,942 fish from 28 species with a total weight of 10,519 kg (23,142 lbs.). Alewife were 89% of the total catch by number while Deepwater Sculpin Myoxocephalus thompsonii , Round Goby Neogobius melanostomus , and Rainbow Smelt Osmerus mordax , comprised 4%, 3%, and 2% of the catch respectively. The estimated Alewife biomass increases slightly from 2023 to 2024 (83.9 to 84.2 kg·ha-1) and was the largest biomass value since whole lake sampling began in 2016. Adult Alewife abundance increased in 2024 as predicted in 2023, and most of the total Alewife biomass was comprised of adult fish (97%), predominantly from the 2020 and 2022 year classes. In contrast, Age-1 Alewife biomass (2.2 kg·ha-1) was the lowest estimated since whole lake sampling began in 2016 (previous range: 2.7 – 26.7 kg·ha-1), indicating reproductive success was poor in 2023. Adult Alewife biomass is predicted to remain relatively high but decline slightly in 2025 and 2026, due to the smaller year classes produced in 2021 and 2023. Alewife condition as measured by the weight of a standard length fish (165 mm; ~6.5 inches), was 32.8 g, which was within of the range of previously observed values (28.0 – 35.9 g, 1997 – 2023). Acoustic-based prey fish densities, in the water above the bottom trawl, were similar to observations from 2021 – 2023 and were orders of magnitude lower than bottom trawl densities. These acoustic results support the seasonal timing of the April survey, when the majority of Alewife and other pelagic prey fishes are near the lake bottom and susceptible to capture with bottom trawls. The trawl survey also provides information on the status of other pelagic prey fishes and native fish restorations. In 2024, biomass indices for Rainbow Smelt, Emerald Shiner Notropis atherinoides , and Threespine Stickleback Gasterosteus aculeatus , were similar to 2023 values while the index for Cisco Coregonus artedi declined. The density index for naturally reproduced, juvenile Lake Trout Salvelinus namaycush declined relative to 2023. Density estimates of Lake Whitefish Coregonus clupeaformis continue to be orders of magnitude lower in U.S. waters relative to Canadian waters. A single purported Bloater Coregonus hoyi (total length = 148 mm, sampling depth = 105 m) was captured near Rochester, NY during the 2024 survey. This is the eighth Bloater recaptured during this survey since restoration stocking began in 2012.

Lake Ontario↗
USGS2023

Larval cisco and lake whitefish exhibit high distributional overlap within nursery areas

Coregonine fishes, including lake whitefish ( Coregonus clupeaformis ) and cisco ( C. artedi ), are socioecologically important in the Laurentian Great Lakes and of conservation concern, but the processes driving recruitment variability are unclear. In Lake Ontario, cisco and lake whitefish exhibit similar spawning behaviours and early life histories, but population trajectories are diverging. One hypothesis is that sympatric cisco and lake whitefish larvae occupy distinct habitats and experience dissimilar local environmental conditions, despite co-occurrence within nursery areas. We described the spatiotemporal distributions of larval cisco and lake whitefish among multiple Lake Ontario embayment nursery areas, characterised physical habitat features associated with their distributions, determined the degree of spatial habitat partitioning between species and evaluated how habitat niche divergence occurred along an ontogenetic progression. Both species were widely distributed across larval nursery areas, though lake whitefish were less abundant and more narrowly distributed than cisco. Within the yolk sac stage, lake whitefish occupied more nearshore, shallower and colder waters than cisco, indicating potential habitat niche partitioning between congeners. However, distributional differences were subtle and likely driven by differential hatch timing and staggered ontogenetic habitat shifts. Combined, our results illustrate similar habitat use between cisco and lake whitefish through the larval stage and demonstrate that ontogeny and species-specific phenology influence habitat use for these species. This study provides additional evidence that the early life histories of cisco and lake whitefish are highly similar and does not support the hypothesis that larval habitat use differences are a major driver of differential recruitment success for these species.

New York↗
USGS2023

Bottom trawl assessment of Lake Ontario's benthic preyfish community, 2022

Since 1978, surveys of Lake Ontario preyfish communities have provided information on the status and trends of the benthic preyfish community related to Fish Community Objectives that includes understanding preyfish population dynamics and community diversity. Beginning in 2015, the benthic preyfish survey expanded from US-only to incorporate Canadian sites, increasing the survey’s spatial coverage to a lake-wide scale. Additionally, sampling in eastern US embayments (Black River, Chaumont, Guffin, and Henderson Bays), that were historically sampled during a September bottom trawl survey to index Yellow Perch ( Perca flavescens ; 1978–2007), resumed in 2015. The current survey provides abundance indices for sculpins, Round Goby ( Neogobius melanostomus ) and Bloater ( Coregonus hoyi ) with survey techniques, gear and timing comparable to Lake Michigan. This alignment provides a necessary biological reference point for measuring the success of Lake Ontario Bloater reintroduction. In 2022, the collaborative benthic preyfish survey completed 171 bottom trawl tows across main lake and embayment sites at depths from 6 to 222 m. In total, the 2022 survey sampled 141,552 fish from 34 species. Round Goby was the most numerically abundant species comprising 36% of the total catch, followed by Alewife ( Alosa pseudoharengus ) and Deepwater Sculpin ( Myoxocephalus thompsonii ), at 20% and 16%, respectively. Alewife accounted for most (623 kg) of the fish biomass sampled during the 2022 survey (total=2,197 kg), followed by Deepwater Sculpin (547 kg), and Round Goby (262 kg). Slimy Sculpin ( Cottus cognatus ) lake-wide biomass density (0.03 kg/ha) remained low relative to historical observations from US waters during the 1980-1990s and was similar to the average from the previous three survey years (2019-2021 average 0.04 ± 0.02 kg/ha). Lake-wide Deepwater Sculpin biomass density reached a new high (4.4 kg/ha) in 2022. Embayment catches continue to have unique species assemblages compared to main lake habitat. Historically common native benthic preyfish species like Trout-perch ( Percopsis omiscomaycus ), Spottail Shiner ( Notropis hudsonius ), and darters ( Etheostoma spp.), that are now rare at main lake trawl sites, still occur in some embayment trawl sites.

Lake Ontario↗
USGS2022

Spatial and temporal variability in lake trout diets in Lake Ontario as revealed by stomach contents and stable isotopes

Lake trout ( Salvelinus namaycush ) are an ecologically and economically important piscivore with reported differences in diet and feeding behaviour throughout its range. Eleven stomach content and stable isotope-based metrics were used to describe diets of 349 lake trout between two years (2013 and 2018) and among geographic zones (west, central, east, Kingston basin) in Lake Ontario. Using individual (e.g., volumetric, %V) and aggregate (e.g., index of relative importance, %IRI) diet metrics, we found an overwhelming dominance of alewife ( Alosa pseudoharengus ) in lake trout diets among some zones in 2013 (%V = 23.3 – 92.7; %IRI = 12.2 – 99.5) and all zones in 2018 (%V = 83.9 – 96.7; %IRI = 96.5 – 100). Round goby ( Neogobius melanostomus ) and rainbow smelt ( Osmerus mordax ) were secondary lake trout prey items with relative diet percentages only marginally reflected by spatial and temporal variation in prey abundance (round goby: %V = 1.0 – 33.3, %IRI = 0.1 – 13.2; rainbow smelt: %V = 2.5 – 54.0, %IRI = 0.1 – 54.0). Carbon (δ 13 C) and nitrogen (δ 15 N) isotopic niche areas and orientations were similar across all year-zone combinations reinforcing temporal and spatial consistency in lake trout diet. The findings of this study advance the time series in describing Lake Ontario lake trout diets and can be used to complement stock assessments and management decisions associated with carrying capacity for the diverse salmonid community.

Lake 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↗
USGS2022

Balancing prey availability and predator consumption: A multispecies stock assessment for Lake Ontario

Trophic interactions are drivers of ecosystem change and stability, yet are often excluded from fishery assessment models, despite their potential capacity to improve estimates of species dynamics and future fishery sustainability. In Lake Ontario, recreational salmonine fisheries, including Chinook salmon ( Oncorhynchus tshawytscha ) and lake trout ( Salvelinus namaycush ), depend on a single prey species, alewife ( Alosa pseudoharengus ). To accommodate strong trophic interactions among species, we developed a multispecies statistical catch-at-age assessment (MSCAA) model that links the dynamics of the salmonine fisheries and alewife via prey consumption and predator growth. We found that prey availability had declined since 2015 due to decreased alewife recruitment and increased Chinook salmon biomass, leading to higher alewife mortality rates and lower predator growth rates. Forward projections of predator–prey dynamics suggest that Chinook salmon stocking reductions may improve the probability for alewife population growth, but could be counteracted by increased natural Chinook salmon recruitment. Combined with predator- and prey-monitoring efforts, multispecies assessments show promise as models of intermediate complexity to support a transition to ecosystem-based approaches to fisheries management.

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

Bottom trawl assessment of benthic preyfish community in Lake Ontario

Since 1978, the Lake Ontario benthic preyfish survey has provided information on the status and trends of the benthic preyfish community related to Lake Ontario Fish Community Objectives that include understanding preyfish population dynamics and community diversity. Beginning in 2015, the benthic preyfish survey expanded from US-only to incorporate lake-wide sampling sites which drastically increased the survey’s spatial coverage. In 2020, the collaborative benthic preyfish survey completed 82 bottom trawl tows across main lake and embayment sites at depths from 6 to 226 m. Compared to previous years, the survey was largely confined to ports east of Rochester on the US side, and east of Wesleyville on the Canadian side due to COVID-19 travel constraints. In total, the survey sampled 109,315 fish from 32 species in 2020. Round goby was the most numerically abundant species comprising 59% of total catch, followed by alewife (27%), and deepwater sculpin (7%). Despite being a common species represented in historical catches, slimy sculpin comprised only 0.05% of the total catch, with only 53 fish sampled in 2020. Annual biomass indices were among the lowest reported for slimy sculpin across the entire time series, whereas deepwater sculpin biomass remained high but showed a decrease from values observed in 2019. Deepwater sculpin condition in 2020 was lower than in 2019 but similar to values observed in 2017-2018. Community diversity remained high relative to historically lower values when round goby and deepwater sculpin were not captured in trawl catches. Round goby biomass in 2020 was the highest on the US time series, but not on the lake-wide series, highlighting the need to resume lake-wide sampling to further understand how regional vs lake-wide sampling affects benthic preyfish population estimates.

Lake Ontario↗
USGS2021

Bottom trawl assessment of Lake Ontario's benthic preyfish community, 2021

Since 1978, the Lake Ontario preyfish community survey has provided information on the status and trends of the benthic preyfish community related to Fish Community Objectives that includes understanding preyfish population dynamics and community diversity. Beginning in 2015, the benthic preyfish survey expanded from US-only to incorporate lake-wide sampling sites which increased the survey’s spatial coverage, and resumed sampling in eastern embayments (Black River, Chaumont, Guffin, and Henderson Bays) that were historically sampled during a September bottom trawl survey to index yellow perch from 1978 to 2007. In 2021, the collaborative benthic preyfish survey completed 195 bottom trawl tows across main lake and embayments at depths from 5 to 226 m. New embayment sites at Bay of Quinte, Sodus, and Little Sodus Bay were added to the survey in 2021 to compare fish communities across nearshore sites. In total, the 2021 survey sampled 109,178 fish from 35 species. Round goby ( Neogobius melanostomus ) was the most numerically abundant species comprising 44% of the total catch, followed by deepwater sculpin ( Myoxocephalus thompsonii ), and alewife ( Alosa pseudoharengus ) at 17% and 11%, respectively. Deepwater sculpin accounted for most (406 kg) of the fish biomass sampled during the 2021 survey (total=1,995 kg), followed by round goby (257 kg), and common carp (252 kg). Slimy sculpin ( Cottus cognatus ) biomass was higher in 2021 than in 2020, when spatial coverage was reduced. Deepwater sculpin biomass remained high in 2021 and similar to observations since 2019. White perch biomass ( Morone americana ) in Black River Bay has increased compared to observations from historical surveys. Yellow perch ( Perca flavescens ) accounted for most of the benthic preyfish biomass across the embayments surveyed in 2021 except for the Bay of Quinte and Black River Bay, where white perch accounted for a greater proportion of the fish community biomass.

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

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↗
USGS2019

Status of Mysis diluviana in Lake Ontario in 2013: lower abundance but higher fecundity than in the 1990s

Mysis diluviana is a major component of prey fish diets in the Great Lakes, so annual production of M. diluviana is important for understanding and modeling energy flow through Great Lakes food webs. However, only three lake-wide measurements of M. diluviana annual production in Lake Ontario are currently available (1971, 1990, 1995). During 2013, lake-wide coverage of Lake Ontario was achieved during four periods from April to November. Annual mean density and biomass of M. diluviana in 2013 were 99 #/m 2 (SE: 8) and 318 mg dw/m 2 (SE: 28) – approximately half of values observed in 1990s. M. diluviana comprised 13–30% of offshore zooplankton biomass in each period. Reproduction peaked in fall, with mean brood size of 32 embryos (range: 11–49), at least 10% larger than in 1990s. Generation time was two years from embryo to initial reproduction. Growth rates were 0.052 mm/d for the age-0 cohort and 0.027 mm/d for the age-1 cohort. Age-0 growth rate was significantly higher than in 1980s–90s (0.035 mm/d). Annual production in 2013 was 0.85 g dw/m 2 /yr (SE: 0.03) which was 30–40% of values observed in 1990 and 1995 (2.23 and 2.53 g/m 2 /yr). Annual production to biomass ratio (P/B) in 2013 was 2.65 /yr which was 80–85% of values observed in 1990 and 1995 (3.24 and 3.11 /yr), but this difference was not statistically significant. Our results suggest that changes in annual production over time can be estimated using changes in biomass over time and a mean P/B ratio.

Lake Ontario↗