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USGS2014

In vivo and in vitro phenotypic differences between Great Lakes VHSV genotype IVb isolates with sequence types vcG001 and vcG002

Viral hemorrhagic septicemia virus (VHSV) is an aquatic rhabdovirus first recognized in farmed rainbow trout in Denmark. In the past decade, a new genotype of this virus, IVb was discovered in the Laurentian Great Lakes basin and has caused several massive die-offs in some of the 28 species of susceptible North American freshwater fishes. Since its colonization of the Great Lakes, several closely related sequence types within genotype IVb have been reported, the two most common of which are vcG001 and vcG002. These sequence types have different spatial distributions in the Great Lakes. The aim of this study was to determine whether the genotypic differences between representative vcG001 (isolate MI03) and vcG002 (isolate 2010-030 #91) isolates correspond to phenotypic differences in terms of virulence using both in vitro and in vivo approaches. In vitro infection of epithelioma papulosum cyprini (EPC), bluegill fry (BF-2), and Chinook salmon embryo (CHSE) cells demonstrated some differences in onset and rate of growth in EPC and BF-2 cells, without any difference in the quantity of RNA produced. In vivo infection of round gobies ( Neogobius melanostomus ) via immersion exposure to different concentrations of vcG001 or vcG002 caused a significantly greater mortality in round gobies exposed to 10 2 plaque forming units ml − 1 of vcG001. These experiments suggest that there are phenotypic differences between Great Lakes isolates of VHSV genotype IVb.

Journal of Great Lakes Research↗

USGS2014

Enterococcus phages as potential tool for identifying sewage inputs in the Great Lakes region

Bacteriophages are viruses living in bacteria that can be used as a tool to detect fecal contamination in surface waters around the world. However, the lack of a universal host strain makes them unsuitable for tracking fecal sources. We evaluated the suitability of two newly isolated Enterococcus host strains (ENT-49 and ENT-55) capable for identifying sewage contamination in impacted waters by targeting phages specific to these hosts. Both host strains were isolated from wastewater samples and identified as E. faecium by 16S rRNA gene sequencing. Occurrence of Enterococcus phages was evaluated in sewage samples ( n = 15) from five wastewater treatment plants and in fecal samples from twenty-two species of wild and domesticated animals (individual samples; n = 22). Levels of Enterococcus phages, F + coliphages, Escherichia coli and enterococci were examined from four rivers, four beaches, and three harbors. Enterococcus phages enumeration was at similar levels (Mean = 6.72 Log PFU/100 mL) to F + coliphages in all wastewater samples, but were absent from all non-human fecal sources tested. The phages infecting Enterococcus spp. and F + coliphages were not detected in the river samples (detection threshold < 10 PFU/100 mL), but were present in the beach and harbor samples (range = 1.83 to 2.86 Log PFU/100 mL). Slightly higher concentrations (range = 3.22 to 3.69 Log MPN/100 mL) of E. coli and enterococci when compared to F + coliphages and Enterococcus phages, were observed in the river, beach and harbor samples. Our findings suggest that the bacteriophages associated with these particular Enterococcus host strains offer potentially sensitive and human-source specific indicators of enteric pathogen risk.

Journal of Great Lakes Research↗

USGS2015

Artificial reefs and reef restoration in the Laurentian Great Lakes

We reviewed the published literature to provide an inventory of Laurentian Great Lakes artificial reef projects and their purposes. We also sought to characterize physical and biological monitoring for artificial reef projects in the Great Lakes and determine the success of artificial reefs in meeting project objectives. We found records of 6 artificial reefs in Lake Erie, 8 in Lake Michigan, 3 in Lakes Huron and Ontario, and 2 in Lake Superior. We found 9 reefs in Great Lakes connecting channels and 6 reefs in Great Lakes tributaries. Objectives of artificial reef creation have included reducing impacts of currents and waves, providing safe harbors, improving sport-fishing opportunities, and enhancing/restoring fish spawning habitats. Most reefs in the lakes themselves were incidental (not created purposely for fish habitat) or built to improve local sport fishing, whereas reefs in tributaries and connecting channels were more frequently built to benefit fish spawning. Levels of assessment of reef performance varied; but long-term monitoring was uncommon as was assessment of physical attributes. Artificial reefs were often successful at attracting recreational species and spawning fish; however, population-level benefits of artificial reefs are unclear. Stressors such as sedimentation and bio-fouling can limit the effectiveness of artificial reefs as spawning enhancement tools. Our investigation underscores the need to develop standard protocols for monitoring the biological and physical attributes of artificial structures. Further, long-term monitoring is needed to assess the benefits of artificial reefs to fish populations and inform future artificial reef projects.

Journal of Great Lakes Research↗

USGS2015

Wildlife, urban inputs, and landscape configuration are responsible for degraded swimming water quality at an embayed beach

Jeorse Park Beach, on southern Lake Michigan, experiences frequent closures due to high Escherichia coli ( E. coli ) levels since regular monitoring was implemented in 2005. During the summer of 2010, contaminant source tracking techniques, such as the conventional microbial and physical surveys and hydrodynamic models, were used to determine the reasons for poor water quality at Jeorse Park. Fecal indicator bacteria ( E. coli , enterococci) were high throughout the season, with densities ranging from 12–2419 (culturable E. coli ) and 1–2550 and < 1–5831 (culturable and qPCR enterococci, respectively). Genetic markers for human ( Bacteroides HF183) and gull ( Catellicoccus marimammalium ) fecal contamination were found in 15% and 37% of the samples indicating multiple sources contributing to poor water quality. Nesting colonies of double-crested cormorants ( Phalacrocorax auritus ) have steadily increased since 2005, coinciding with high E. coli levels. A hydrodynamic model indicated that limited circulation allows bacteria entering the embayed area to be retained in nearshore areas; and bacterial resuspension from sand and stranded beach wrack during storm events compounds the problem. The integration of hydrodynamics, expanded use of chemical and biological markers, as well as more complex statistical multivariate techniques can improve microbial source tracking, informing management actions to improve recreational water quality. Alterations to embayed structures to improve circulation and reduce nuisance algae as well as growing native plants to retain sand to improve beach morphometry are among some of the restoration strategies under consideration in ongoing multi-agency collaborations.

Indiana↗

USGS2015

Non-stationary recruitment dynamics of rainbow smelt: the influence of environmental variables and variation in size structure and length-at-maturation

Fish stock-recruitment dynamics may be difficult to elucidate because of nonstationary relationships resulting from shifting environmental conditions and fluctuations in important vital rates such as individual growth or maturation. The Great Lakes have experienced environmental stressors that may have changed population demographics and stock-recruitment relationships while causing the declines of several prey fish species, including rainbow smelt ( Osmerus mordax ). We investigated changes in the size and maturation of rainbow smelt in Lake Michigan and Lake Huron and recruitment dynamics of the Lake Michigan stock over the past four decades. Mean lengths and length-at-maturation of rainbow smelt generally declined over time in both lakes. To evaluate recruitment, we used both a Ricker model and a Kalman filter-random walk (KF-RW) model which incorporated nonstationarity in stock productivity by allowing the productivity term to vary over time. The KF-RW model explained nearly four times more variation in recruitment than the Ricker model, indicating the productivity of the Lake Michigan stock has increased. By accounting for this nonstationarity, we were able identify significant variations in stock productivity, evaluate its importance to rainbow smelt recruitment, and speculate on potential environmental causes for the shift. Our results suggest that investigating mechanisms driving nonstationary shifts in stock-recruit relationships can provide valuable insights into temporal variation in fish population dynamics.

Lake Huron, Lake Michigan↗

USGS2015

From yellow perch to round goby: A review of double-crested cormorant diet and fish consumption at three St. Lawrence River colonies, 1999–2013

The number of double-crested cormorants ( Phalacrocorax auritus ) in the upper St. Lawrence River has increased markedly since the early 1990s. In 1999, a binational study was initiated to examine the annual diet composition and fish consumption of cormorants at colonies in the upper river. Since 1999, 14,032 cormorant pellets, collected from May through September each year, have been examined from St. Lawrence River colonies to estimate fish consumption and determine temporal and spatial variation in diet. Seasonal variation in diet composition within a colony was low. Prior to 2006 yellow perch was the primary fish consumed by cormorants in the upper St. Lawrence River. Round goby were first observed in cormorant diets in 2003 and by 2006 were the main fish consumed at two of the three colonies. The time interval it took from the first appearance of round goby in the diet at a colony to when goby were the dominant prey species varied by island, ranging from two to five years. Daily fish consumption at each cormorant colony increased significantly from the pre-round goby to post-round goby period. The mean annual biomass of yellow perch consumed decreased significantly during the post-round goby period at the three colonies. Reduced consumption of yellow perch by cormorants may alleviate suspected localized impacts on perch near some of the larger river colonies.

New York, Ontario↗

USGS2015

Sea lamprey mark type, marking rate, and parasite-host relationships for lake trout and other species in Lake Ontario

We examined how attack frequency by sea lampreys on fishes in Lake Ontario varied in response to sea lamprey abundance and preferred host abundance (lake trout > 433 mm). For this analysis we used two gill net assessment surveys, one angler creel survey, three salmonid spawning run datasets, one adult sea lamprey assessment, and a bottom trawl assessment of dead lake trout. The frequency of fresh sea lamprey marks observed on lake trout from assessment surveys was strongly related to the frequency of sea lamprey attacks observed on salmon and trout from the creel survey and spawning migrations. Attack frequencies on all salmonids examined were related to the ratio between the abundances of adult sea lampreys and lake trout. Reanalysis of the susceptibility to sea lamprey attack for lake trout strains stocked into Lake Ontario reaffirmed that Lake Superior strain lake trout were among the most and Seneca Lake strain among the least susceptible and that Lewis Lake strain lake trout were even more susceptible than the Superior strain. Seasonal attack frequencies indicated that as the number of observed sea lamprey attacks decreased during June–September, the ratio of healing to fresh marks also decreased. Simulation of the ratios of healing to fresh marks indicated that increased lethality of attacks by growing sea lampreys contributed to the decline in the ratios and supported laboratory studies about wound healing duration.

Lake Ontario↗

USGS2015

Application of the FluEgg model to predict transport of Asian carp eggs in the Saint Joseph River (Great Lakes tributary)

The Fluvial Egg Drift Simulator (FluEgg) is a three-dimensional Lagrangian model that simulates the movement and development of Asian carp eggs until hatching based on the physical characteristics of the flow field and the physical and biological characteristics of the eggs. This tool provides information concerning egg development and spawning habitat suitability including: egg plume location, egg vertical and travel time distribution, and egg-hatching risk. A case study of the simulation of Asian carp eggs in the Lower Saint Joseph River, a tributary of Lake Michigan, is presented. The river hydrodynamic input for FluEgg was generated in two ways — using hydroacoustic data and using HEC-RAS model data. The HEC-RAS model hydrodynamic input data were used to simulate 52 scenarios covering a broad range of flows and water temperatures with the eggs at risk of hatching ranging from 0 to 93% depending on river conditions. FluEgg simulations depict the highest percentage of eggs at risk of hatching occurs at the lowest discharge and at peak water temperatures. Analysis of these scenarios illustrates how the interactive relation among river length, hydrodynamics, and water temperature influence egg transport and hatching risk. An improved version of FluEgg, which more realistically simulates dispersion and egg development, is presented. Also presented is a graphical user interface that facilitates the use of FluEgg and provides a set of post-processing analysis tools to support management decision-making regarding the prevention and control of Asian carp reproduction in rivers with or without Asian carp populations.

Michigan↗

USGS2015

Potential impact of Chironomus plumosus larvae on hypolimnetic oxygen in the central basin of Lake Erie

Previous studies have indicated that burrow-irrigating infauna can increase sediment oxygen demand (SOD) and impact hypolimnetic oxygen in stratified lakes. We conducted laboratory microcosm experiments and computer simulations with larvae of the burrowing benthic midge Chironomus plumosus to quantify burrow oxygen uptake rates and subsequent contribution to sediment oxygen demand in central Lake Erie. Burrow oxygen uptake and water flow velocities through burrows were measured using oxygen microelectrodes and hot film anemometry, respectively. Burrow oxygen consumption averaged 2.66 × 10 − 10 (SE = ± 7.82 × 10 − 11 ) mol O 2 /burrow/s at 24 °C and 9.64 × 10 − 10 (SE = ± 4.86 × 10 − 10 ) mol O 2 /burrow/s at 15 °C. In sealed microcosm experiments, larvae increased SOD 500% at 24 °C (density = 1508/m 2 ) and 375% at 15 °C (density = 864/m 2 ). To further evaluate effects of densities of C. plumosus burrows on SOD we developed a 3-D transport reaction model of the process. Using experimental data and chironomid abundance data in faunal surveys in 2009 and 2010, we estimated that bioirrigation by a population of 140 larvae/m 2 could account for between 2.54 × 10 − 11 mol/L/s (model results) and 5.58 × 10 − 11 mol/L/s (experimental results) of the average 4.22 × 10 − 11 mol/L/s oxygen depletion rate between 1970 and 2003, which could have accounted for 60–132% of the oxygen decline. At present, it appears that the population density of this species may be an important factor in development of hypoxic or anoxic conditions in central Lake Erie.

Ohio↗

USGS2015

Improving efficiency and reliability of environmental DNA analysis for silver carp

Natural resource agencies have established surveillance programs which use environmental DNA (eDNA) for the early detection of bighead carp Hypophthalmichthys nobilis and silver carp Hypophthalmichthys molitrix before they establish populations within the Great Lakes. This molecular monitoring technique must be highly accurate and precise for confident interpretation and also efficient, both in detection threshold and cost. Therefore, we compared two DNA extraction techniques and compared a new quantitative PCR (qPCR) assay with the conventional PCR (cPCR) assay used by monitoring programs. Both the qPCR and cPCR assays were able to amplify the DNA of silver carp present in environmental samples taken from locations where mixed populations of bigheaded carps existed. However, the qPCR assay had substantially fewer PCR positive samples which were subsequently determined not to contain DNA of bigheaded carps than the cPCR assay. Additionally, the qPCR assay was able to amplify the DNA of bigheaded carps even in the presence of inhibitors that blocked amplification with cPCR. Also, the selection of an appropriate DNA extraction method can significantly alter the efficiency of eDNA surveillance programs by lowering detection limits and by decreasing costs associated with sample processing. The results reported herein are presently being incorporated into eDNA surveillance programs to decrease the costs, increase DNA yield and increase the confidence that assays are amplifying the target DNA. These results are critical to enhancing our ability to accurately and confidently interpret the results reported from monitoring programs using eDNA for early detection of invasive species.

Journal of Great Lakes Research↗

USGS2015

Natural trophic variability in a large, oligotrophic, near-pristine lake

Conclusions drawn from stable isotope data can be limited by an incomplete understanding of natural isotopic variability over time and space. We quantified spatial and temporal variability in fish carbon and nitrogen stable isotopes in Lake Hövsgöl, Mongolia, a large, remote, oligotrophic lake with an unusually species-poor fish community. The fish community demonstrated a high degree of trophic level overlap. Variability in δ 13 C was inversely related to littoral-benthic dependence, with pelagic species demonstrating more δ 13 C variability than littoral-benthic species. A mixed effects model suggested that space (sampling location) had a greater impact than time (collection year) on both δ 13 C and δ 15 N variability. The observed variability in Lake Hövsgöl was generally greater than isotopic variability documented in other large, oligotrophic lakes, similar to isotopic shifts attributed to introduced species, and less than isotopic shifts attributed to anthropogenic chemical changes such as eutrophication. This work complements studies on isotopic variability and changes in other lakes around the world.

Journal of Great Lakes Research↗

USGS2015

Potential effects of climate change on the growth of fishes from different thermal guilds in Lakes Michigan and Huron

We used a bioenergetics modeling approach to investigate potential effects of climate change on the growth of two economically important native fishes: yellow perch ( Perca flavescens ), a cool-water fish, and lake whitefish ( Coregonus clupeaformis ), a cold-water fish, in deep and oligotrophic Lakes Michigan and Huron. For assessing potential changes in fish growth, we contrasted simulated fish growth in the projected future climate regime during the period 2043-2070 under different prey availability scenarios with the simulated growth during the baseline (historical reference) period 1964-1993. Results showed that effects of climate change on the growth of these two fishes are jointly controlled by behavioral thermoregulation and prey availability. With the ability of behavioral thermoregulation, temperatures experienced by yellow perch in the projected future climate regime increased more than those experienced by lake whitefish. Thus simulated future growth decreased more for yellow perch than for lake whitefish under scenarios where prey availability remains constant into the future. Under high prey availability scenarios, simulated future growth of these two fishes both increased but yellow perch could not maintain the baseline efficiency of converting prey consumption into body weight. We contended that thermal guild should not be the only factor used to predict effects of climate change on the growth of a fish, and that ecosystem responses to climate change should be also taken into account.

Lake Huron, Lake Michigan↗

USGS2015

Avian botulism type E in waterbirds of Lake Michigan, 2010–2013

During 2010 to 2013, waterbird mortality surveillance programs used a shared protocol for shoreline walking surveys performed June to November at three areas in northern Lake Michigan. In 2010 and 2012, 1244 total carcasses (0.8 dead bird/km walked) and 2399 total carcasses (1.2 dead birds/km walked), respectively, were detected. Fewer carcasses were detected in 2011 (353 total carcasses, 0.2 dead bird/km walked) and 2013 (451 total carcasses, 0.3 dead bird/km walked). During 3 years, peak detection of carcasses occurred in October and involved primarily migratory diving and fish-eating birds, including long-tailed ducks ( Clangula hyemalis ; 2010), common loons ( Gavia immer ; 2012), and red-breasted mergansers ( Mergus serrator ; 2013). In 2011, peak detection of carcasses occurred in August and consisted primarily of summer residents such as gulls ( Larus spp.) and double-crested cormorants ( Phalacrocorax auritus ). A subset of fresh carcasses was collected throughout each year of the study and tested for botulinum neurotoxin type E (BoNT/E). Sixty-one percent of carcasses (57/94) and 10 of 11 species collected throughout the sampling season tested positive for BoNT/E, suggesting avian botulism type E was a major cause of death for both resident and migratory birds in Lake Michigan. The variety of avian species affected by botulism type E throughout the summer and fall during all 4 years of coordinated surveillance also suggests multiple routes for bird exposure to BoNT/E in Lake Michigan.

Lake Michigan↗

USGS2015

Thiaminase activity in native freshwater mussels

Thiamine (vitamin B 1 ) deficiency in the Great Lakes has been attributed to elevated levels of thiaminase I enzyme activity in invasive prey species; however, few studies have investigated thiaminase activity in native prey species. Some of the highest levels of thiaminase activity have been measured in invasive dreissenid mussels with little understanding of background levels contributed by native freshwater mussels (Bivalvia: Unionidae). In this study, thiaminase activity was measured in two freshwater mussel species, Elliptio complanata and Strophitus undulatus , from the Delaware and Susquehanna River drainage basins located in north eastern United States. Thiaminase activity was also measured in gravid and non-gravid S. undulatus . Average thiaminase activity differed significantly between species (7.2 and 42.4 μmol/g/min, for E. complanata and S. undulatus respectively) with no differences observed between drainage basins. Gravid S. undulatus had significantly lower thiaminase activity (28.0 μmol/g/min) than non-gravid mussels (42.4 μmol/g/min). Our results suggest that a suite of factors may regulate thiaminase activity in freshwater mussels and that native freshwater mussel thiaminase activity is within the range observed for invasive dreissenids. These results add to our understanding of the complexities in identifying the ecological conditions that set the stage for thiamine deficiency.

Journal of Great Lakes Research↗

USGS2015

Suitability of Lake Erie for bigheaded carps based on bioenergetic models and remote sensing

Algal blooms in the Great Lakes are a potential food source for silver carp ( Hypophthalmichthys molitrix ) and bighead carp ( H. nobilis ; together bigheaded carps). Understanding these blooms thus plays an important role in understanding the invasion potential of bigheaded carps. We used remote sensing imagery, temperatures, and improved species specific bioenergetics models to determine algal concentrations sufficient for adult bigheaded carps. Depending on water temperature we found that bigheaded carp require between 2 and 7 μg/L chlorophyll or between 0.3 and 1.26 × 10 5 cells/mL Microcystis to maintain body weight. Algal concentrations in the western basin and shoreline were found to be commonly several times greater than the concentrations required for weight maintenance. The remote sensing images show that area of sufficient algal foods commonly encompassed several hundred square kilometers to several thousands of square kilometers when blooms form. From 2002 to 2011, mean algal concentrations increased 273%–411%. This indicates Lake Erie provides increasingly adequate planktonic algal food for bigheaded carps. The water temperatures and algal concentrations detected in Lake Erie from 2008 to 2012 support positive growth rates such that a 4 kg silver carp could gain between 19 and 57% of its body weight in a year. A 5 kg bighead carp modeled at the same water temperatures could gain 20–81% of their body weight in the same period. The remote sensing imagery and bioenergetic models suggest that bigheaded carps would not be food limited if they invaded Lake Erie.

Lake Erie, Lake St. Clair↗

USGS2015

Little Galloo Island, Lake Ontario: Two decades of studies on the diet, fish consumption, and management of double-crested cormorants

The double-crested cormorant ( Phalacrocorax auritus ) colony at Little Galloo Island, Lake Ontario has been a Great Lakes focal point of controversy regarding cormorant–fish interactions for over two decades. We examined cormorant diet and fish consumption at the colony from 1992 to 2013. During this time period, two events, management actions and round goby ( Neogobius melanostomus ) invasion, occurred that affected the number of fish consumed by cormorants and their diet composition. The purpose of this study was to evaluate the effects of round goby on the feeding ecology of cormorants and evaluate the efficacy of management actions on meeting cormorant population targets at the colony. Round goby first appeared in the diet in 2004 (0.8%) and within one year were the primary prey (29.3%). The presence of round goby in the diet of cormorants: (1) eliminated seasonal variability in diet composition, (2) reversed seasonal trends in the number of fish consumed daily, (3) increased daily fish consumption, and (4) significantly reduced the consumption of other species including yellow perch and smallmouth bass. Management actions, such as egg oiling and culling, were also effective in reducing nesting activity and the number of cormorant feeding days at the Little Galloo Island colony. There is evidence that the combination of management actions and round goby may have allowed some population recovery of yellow perch and smallmouth bass in eastern Lake Ontario.

New York↗

USGS2015

Stock-specific advection of larval walleye ( Sander vitreus ) in western Lake Erie: Implications for larval growth, mixing, and stock discrimination

Physical processes can generate spatiotemporal heterogeneity in habitat quality for fish and also influence the overlap of pre-recruit individuals (e.g., larvae) with high-quality habitat through hydrodynamic advection. In turn, individuals from different stocks that are produced in different spawning locations or at different times may experience dissimilar habitat conditions, which can underlie within- and among-stock variability in larval growth and survival. While such physically-mediated variation has been shown to be important in driving intra- and inter-annual patterns in recruitment in marine ecosystems, its role in governing larval advection, growth, survival, and recruitment has received less attention in large lake ecosystems such as the Laurentian Great Lakes. Herein, we used a hydrodynamic model linked to a larval walleye ( Sander vitreus ) individual-based model to explore how the timing and location of larval walleye emergence from several spawning sites in western Lake Erie (Maumee, Sandusky, and Detroit rivers; Ohio reef complex) can influence advection pathways and mixing among these local spawning populations (stocks), and how spatiotemporal variation in thermal habitat can influence stock-specific larval growth. While basin-wide advection patterns were fairly similar during 2011 and 2012, smaller scale advection patterns and the degree of stock mixing varied both within and between years. Additionally, differences in larval growth were evident among stocks and among cohorts within stocks which were attributed to spatiotemporal differences in water temperature. Using these findings, we discuss the value of linked physical–biological models for understanding the recruitment process and addressing fisheries management problems in the world's Great Lakes.

Journal of Great Lakes Research↗

USGS2015

Evaluating the importance of abiotic and biotic drivers on Bythotrephes biomass in Lakes Superior and Michigan

The ability of planktivorous fishes to exert top-down control on Bythotrephes potentially has far-reaching impacts on aquatic food-webs, given previously described effects of Bythotrephes on zooplankton communities. We estimated consumption of Bythotrephes by planktivorous and benthivorous fishes, using bioenergetics and daily ration models at nearshore (18 m), intermediate (46 m), and offshore (110 m) depths along one western Lake Superior transect (April, and September-November) and two northern Lake Michigan transects (April, July, September). In Lake Superior, consumption (primarily by cisco Coregonus artedi) exceeded Bythotrephes production at all offshore sites in September-November (up to 396% of production consumed) and at the intermediate site in November (842%) with no evidence of consumption nearshore. By comparing Bythotrephes biomass following months of excessive consumption, we conservatively concluded that top-down control was evident only at the offshore site during September-October. In Lake Michigan, consumption by fishes (primarily alewife Alosa pseudoharengus) exceeded production at nearshore sites (up to 178%), but not in deeper sites (< 15%). Evidence for top-down control in the nearshore was not supported, however, as Bythotrephes never subsequently declined. Using generalized additive models, temperature, and not fish consumption, not zooplankton prey density, best explained variability in Bythotrephes biomass. The non-linear pattern revealed Bythotrephes to increase with temperature up to 16 °C, and then decline between 16 and 23 °C. We discuss how temperature likely has direct negative impacts on Bythotrephes when temperatures near 23 °C, but speculate that predation also contributes to declining biomass when temperatures exceed 16 °C.

Wisconsin and Michigan↗