Potential changes to the biology and challenges to the management of invasive sea lamprey Petromyzon marinus in the Laurentian Great Lakes due to climate change.

Control programs are implemented to mitigate the damage caused by invasive species worldwide. In the highly invaded Great Lakes, the climate is expected to become warmer with more extreme weather and variable precipitation, resulting in shorter iced-over periods and variable tributary flows as well as changes to pH and river hydrology and hydrogeomorphology. We review how climate change influences physiology, behavior, and demography of a damaging invasive species, sea lamprey (Petromyzon marinus), in the Great Lakes, and the consequences for sea lamprey control efforts. Sea lamprey control relies on surveys to monitor abundance of larval sea lamprey in Great Lakes tributaries. The abundance of parasitic, juvenile sea lampreys in the lakes is calculated by surveying wounding rates on lake trout (Salvelinus namaycush), and trap surveys are used to enumerate adult spawning runs. Chemical control using lampricides (i.e., lamprey pesticides) to target larval sea lamprey and barriers to prevent adult lamprey from reaching spawning grounds are the most important tools used for sea lamprey population control. We describe how climate change could affect larval survival in rivers, growth and maturation in lakes, phenology and the spawning migration as adults return to rivers, and the overall abundance and distribution of sea lamprey in the Great Lakes. Our review suggests that Great Lakes sea lamprey may benefit from climate change with longer growing seasons, more rapid growth, and greater access to spawning habitat, but uncertainties remain about the future availability and suitability of larval habitats. Consideration of the biology of invasive species and adaptation of the timing, intensity, and frequency of control efforts is critical to the management of biological invasions in a changing world, such as sea lamprey in the Great Lakes.

Absence of PCB Hot Spot Effect in Walleye Sander vitreus from Lower Green Bay of Lake Michigan.

Under certain conditions, polychlorinated biphenyl (PCB) concentration in individuals of one sex of an adult fish population may exceed that of the other sex by more than a factor of two. This phenomenon, known as the PCB hot spot effect, has been postulated to be contingent upon the following two conditions: (1) presence of a PCB hot spot in the bottom sediments of the aquatic ecosystem, such that prey PCB concentrations in the hot spot region are substantially higher than prey PCB concentrations in locations distant from the hot spot, and (2) habitat use varying between the sexes, such that individuals of one sex inhabit the hot spot region to a considerably greater degree than individuals of the other sex. To test whether PCB concentrations in walleye Sander vitreus from lower Green Bay of Lake Michigan displayed a PCB hot spot effect, whole-fish PCB concentrations were determined in ten female and ten male adult walleye from the population spawning in the Fox River, the main tributary to lower Green Bay. In addition, mark-recapture data for the Fox River walleye population were analyzed to determine differences in spatial distributions between the sexes. Results revealed that the ratio of mean PCB concentration in males to mean PCB concentration in females was only 1.13, indicating the absence of a PCB hot spot effect. This result was corroborated by the mark-recapture data analysis, which showed no significant difference in habitat use between the sexes. Thus, although condition 1 was met, condition 2 was not met. Consequently, the PCB hot spot effect was not observed in the Fox River walleye population. Lack of a significant difference in PCB congener distributions between the sexes further corroborated our conclusions.

Distributions of PCB Congeners and Homologues in White Sucker and Coho Salmon from Lake Michigan.

We tested the hypothesis of the proportion of higher chlorinated biphenyl (PCB) congeners increasing with increasing trophic level by comparing the respective PCB homologue distributions in an omnivore, white sucker ( Catostomus commersoni), and a top predator, coho salmon ( Oncorhynchus kisutch), from Lake Michigan. Adult females had the same congener and homologue proportions of total PCB concentration (ΣPCB) as adult males in both species. Hexachlorinated congeners comprised the largest proportion (32%) found in white sucker, followed by heptachlorinated (21%) and octochlorinated (18%) congeners. In contrast, pentachlorinated congeners comprised the largest proportion (33%) of ΣPCB found in coho salmon, followed by hexachlorinated (26%) and tetrachlorinated (24%) congeners. Coho salmon contained significantly higher proportions of tri-, tetra-, and pentachlorinated congeners, whereas white sucker contained significantly higher proportions of hexa- through decachlorinated congeners. Our results were opposite of the hypothesis of greater degree of PCB chlorination with increasing trophic level, and supported the contention that the PCB congener proportions in fish depends mainly on diet, and does not necessarily reflect the trophic level of the fish. Our results also supported the contention that diets do not vary between the sexes in most fish populations.

Females exceed males in mercury concentrations of burbot Lota lota.

Examination of differences in contaminant concentrations between the sexes of fish, across several fish species, may show clues for important behavioral and physiological differences between the sexes. We determined whole-fish total mercury (Hg) concentrations of 25 male and 25 female adult burbot Lota lota captured in Lake Erie during summer 2011 and of 14 male and 18 female adult burbot captured in Great Slave Lake (Northwest Territories, Canada) during winter 2013. On average, females had 22 % greater Hg concentrations than males. This difference was probably not due to a greater feeding rate by females because results from previous studies based on polychlorinated biphenyl determinations of these same burbot indicated that males fed at a substantially greater rate than females. Based on our determinations of Hg concentrations in the gonads and somatic tissue of 5 ripe females and 5 ripe males, this difference was not attributable to changes in Hg concentration immediately after spawning due to the release of gametes. Furthermore, bioenergetics modeling results from previous studies indicated that growth dilution would not explain any portion of this observed difference in Hg concentrations between the sexes. We therefore conclude that this difference was most likely due to a substantially faster rate of Hg elimination by males compared with females. Male burbot exhibit among the greatest gonadosomatic indices (GSIs) of all male fishes, with their testes accounting for between 10 and 15 % of their body weight when the fish are in ripe condition. Androgens have been linked to enhanced Hg-elimination rates in other vertebrates. If androgen production is positively related to GSI, then male burbot would be expected to have among the greatest androgen levels of all fishes. Thus, we hypothesize that male burbot eliminate Hg from their bodies faster than most other male fishes and that this explains the greater Hg concentration in females compared with males.

Seasonal thermal ecology of adult walleye (Sander vitreus) in Lake Huron and Lake Erie.

The purpose of this study was to characterize thermal patterns and generate occupancy models for adult walleye from lakes Erie and Huron with internally implanted biologgers coupled with a telemetry study to assess the effects of sex, fish size, diel periods, and lake. Sex, size, and diel periods had no effect on thermal occupancy of adult walleye in either lake. Thermal occupancy differed between lakes and seasons. Walleye from Lake Erie generally experienced higher temperatures throughout the spring and summer months than did walleye in Lake Huron, due to limnological differences between the lakes. Tagged walleye that remained in Saginaw Bay, Lake Huron (i.e., adjacent to the release location), as opposed to those migrating to the main basin of Lake Huron, experienced higher temperatures, and thus accumulated more thermal units (the amount of temperature units amassed over time) throughout the year. Walleye that migrated toward the southern end of Lake Huron occupied higher temperatures than those that moved toward the north. Consequently, walleye that emigrated from Saginaw Bay experienced thermal environments that were more favorable for growth as they spent more time within their thermal optimas than those that remained in Saginaw Bay. Results presented in this paper provide information on the thermal experience of wild fish in a large lake, and could be used to refine sex- and lake-specific bioenergetics models of walleye in the Great Lakes to enable the testing of ecological hypotheses.

Mercury elimination rates for adult northern pike Esox lucius: evidence for a sex effect.

We examined the effect of sex on mercury elimination in fish by monitoring isotope-enriched mercury concentrations in the muscle tissue of three adult female and three adult male northern pike Esox lucius, which had accumulated the isotope-enriched mercury via a whole-lake manipulation and were subsequently moved to a clean lake. Mercury elimination rates for female and male northern pike were estimated to be 0.00034 and 0.00073 day(-1), respectively. Thus, males were capable of eliminating mercury at more than double the rate than that of females. To the best of our knowledge, our study represents the first documentation of mercury elimination rates varying between the sexes of fish. This sex difference in elimination rates should be taken into account when comparing mercury accumulation between the sexes of fish from the same population. Further, our findings should eventually lead to an improved understanding of mechanisms responsible for mercury elimination in vertebrates.

Reevaluation of a walleye (Sander vitreus) bioenergetics model.

Walleye (Sander vitreus) is an important sport fish throughout much of North America, and walleye populations support valuable commercial fisheries in certain lakes as well. Using a corrected algorithm for balancing the energy budget, we reevaluated the performance of the Wisconsin bioenergetics model for walleye in the laboratory. Walleyes were fed rainbow smelt (Osmerus mordax) in four laboratory tanks each day during a 126-day experiment. Feeding rates ranged from 1.4 to 1.7% of walleye body weight per day. Based on a statistical comparison of bioenergetics model predictions of monthly consumption with observed monthly consumption, we concluded that the bioenergetics model estimated food consumption by walleye without any significant bias. Similarly, based on a statistical comparison of bioenergetics model predictions of weight at the end of the monthly test period with observed weight, we concluded that the bioenergetics model predicted walleye growth without any detectable bias. In addition, the bioenergetics model predictions of cumulative consumption over the 126-day experiment differed from observed cumulative consumption by less than 10%. Although additional laboratory and field testing will be needed to fully evaluate model performance, based on our laboratory results, the Wisconsin bioenergetics model for walleye appears to be providing unbiased predictions of food consumption.

Absence of fractionation of mercury isotopes during trophic transfer of methylmercury to freshwater fish in captivity.

We performed two controlled experiments to determine the amount of mass-dependent and mass-independent fractionation (MDF and MIF) of methylmercury (MeHg) during trophic transfer into fish. In experiment 1, juvenile yellow perch (Perca flavescens) were raised in captivity on commercial food pellets and then their diet was either maintained on unamended food pellets (0.1 µg/g MeHg) or was switched to food pellets with 1.0 µg/g or 4.0 µg/g of added MeHg, for a period of 2 months. The difference in δ(202)Hg (MDF) and Δ(199)Hg (MIF) between fish tissues and food pellets with added MeHg was within the analytical uncertainty (δ(202)Hg, 0.07 ‰; Δ(199)Hg, 0.06 ‰), indicating no isotope fractionation. In experiment 2, lake trout (Salvelinus namaycush) were raised in captivity on food pellets and then shifted to a diet of bloater (Coregonus hoyi) for 6 months. The δ(202)Hg and Δ(199)Hg of the lake trout equaled the isotopic composition of the bloater after 6 months, reflecting reequilibration of the Hg isotopic composition of the fish to new food sources and a lack of isotope fractionation during trophic transfer. We suggest that the stable Hg isotope ratios in fish can be used to trace environmental sources of Hg in aquatic ecosystems.

Population synchrony of a native fish across three Laurentian Great Lakes: evaluating the effects of dispersal and climate.

Climate and dispersal are the two most commonly cited mechanisms to explain spatial synchrony among time series of animal populations, and climate is typically most important for fishes. Using data from 1978-2006, we quantified the spatial synchrony in recruitment and population catch-per-unit-effort (CPUE) for bloater (Coregonus hoyi) populations across lakes Superior, Michigan, and Huron. In this natural field experiment, climate was highly synchronous across lakes but the likelihood of dispersal between lakes differed. When data from all lakes were pooled, modified correlograms revealed spatial synchrony to occur up to 800 km for long-term (data not detrended) trends and up to 600 km for short-term (data detrended by the annual rate of change) trends. This large spatial synchrony more than doubles the scale previously observed in freshwater fish populations, and exceeds the scale found in most marine or estuarine populations. When analyzing the data separately for within- and between-lake pairs, spatial synchrony was always observed within lakes, up to 400 or 600 km. Conversely, between-lake synchrony did not occur among short-term trends, and for long-term trends, the scale of synchrony was highly variable. For recruit CPUE, synchrony occurred up to 600 km between both lakes Michigan and Huron (where dispersal was most likely) and lakes Michigan and Superior (where dispersal was least likely), but failed to occur between lakes Huron and Superior (where dispersal likelihood was intermediate). When considering the scale of putative bloater dispersal and genetic information from previous studies, we concluded that dispersal was likely underlying within-lake synchrony but climate was more likely underlying between-lake synchrony. The broad scale of synchrony in Great Lakes bloater populations increases their probability of extirpation, a timely message for fishery managers given current low levels of bloater abundance.

Laboratory evaluation of a walleye (Sander vitreus) bioenergetics model.

Walleye (Sander vitreus) is an important game fish throughout much of North America. We evaluated the performance of the Wisconsin bioenergetics model for walleye in the laboratory. Walleyes were fed rainbow smelt (Osmerus mordax) in four laboratory tanks during a 126-day experiment. Based on a statistical comparison of bioenergetics model predictions of monthly consumption with the observed monthly consumption, we concluded that the bioenergetics model significantly underestimated food consumption by walleye in the laboratory. The degree of underestimation appeared to depend on the feeding rate. For the tank with the lowest feeding rate (1.4% of walleye body weight per day), the agreement between the bioenergetics model prediction of cumulative consumption over the entire 126-day experiment and the observed cumulative consumption was remarkably close, as the prediction was within 0.1% of the observed cumulative consumption. Feeding rates in the other three tanks ranged from 1.6% to 1.7% of walleye body weight per day, and bioenergetics model predictions of cumulative consumption over the 126-day experiment ranged between 11 and 15% less than the observed cumulative consumption.

Restoring piscivorous fish populations in the Laurentian Great Lakes causes seabird dietary change.

Ecosystem change often affects the structure of aquatic communities thereby regulating how much and by what pathways energy and critical nutrients flow through food webs. The availability of energy and essential nutrients to top predators such as seabirds that rely on resources near the water's surface will be affected by changes in pelagic prey abundance. Here, we present results from analysis of a 25-year data set documenting dietary change in a predatory seabird from the Laurentian Great Lakes. We reveal significant declines in trophic position and alterations in energy and nutrient flow over time. Temporal changes in seabird diet tracked decreases in pelagic prey fish abundance. As pelagic prey abundance declined, birds consumed less aquatic prey and more terrestrial food. This pattern was consistent across all five large lake ecosystems. Declines in prey fish abundance may have primarily been the result of predation by stocked piscivorous fishes, but other lake-specific factors were likely also important. Natural resource management activities can have unintended consequences for nontarget ecosystem components. Reductions in pelagic prey abundance have reduced the capacity of the Great Lakes to support the energetic requirements of surface-feeding seabirds. In an environment characterized by increasingly limited pelagic fish resources, they are being offered a Hobsonian choice: switch to less nutritious terrestrial prey or go hungry.

Net trophic transfer efficiencies of polychlorinated biphenyl congeners to lake whitefish (Coregonus clupeaformis) from their food.

Lake whitefish (Coregonus clupeaformis) were fed rainbow smelt (Osmerus mordax) in four laboratory tanks over a 133-d experiment. At the start of the experiment, 10 to 14 of the fish in each tank were sacrificed, and the concentrations of 40 polychlorinated biphenyl (PCB) congeners within these fish were determined. Polychlorinated biphenyl congener concentrations were also determined in the 15 lake whitefish remaining in each of the four tanks at the end of the experiment as well as in the rainbow smelt fed to the lake whitefish. Each lake whitefish was weighed at the start and the end of the experiment, and the amount of food eaten by the lake whitefish during the experiment was tracked. Using these measurements, net trophic transfer efficiency (gamma) from the rainbow smelt to the lake whitefish in each of the four tanks was calculated for each of the 40 PCB congeners. Results showed that gamma decreased exponentially as log K(OW) for the congeners increased from 6 to 8. Further, gamma averaged 0.70 for the tetrachloro congeners but averaged only 0.45 for the higher chlorinated congeners.

Differences in Energy Expenditures and Growth Dilution Explain Higher PCB Concentrations in Male Summer Flounder.

Comparison of polychlorinated biphenyl (PCB) concentrations between the sexes of mature fish may reveal important behavioral and physiological differences between the sexes. We determined whole-fish PCB concentrations in 23 female summer flounder Paralichthys dentatus and 27 male summer flounder from New Jersey coastal waters. To investigate the potential for differences in diet or habitat utilization between the sexes, carbon and nitrogen stable isotope ratios were also determined. In 5 of the 23 female summer flounder, PCB concentrations in the somatic tissue and ovaries were determined. In addition, we used bioenergetics modeling to assess the contribution of the growth dilution effect to the observed difference in PCB concentrations between the sexes. Whole-fish PCB concentrations for females and males averaged 87 and 124 ng/g, respectively; thus males were 43% higher in PCB concentration compared with females. Carbon and nitrogen stable isotope ratios did not significantly differ between the sexes, suggesting that diet composition and habitat utilization did not vary between the sexes. Based on PCB determinations in the somatic tissue and ovaries, we predicted that PCB concentration of females would increase by 0.6%, on average, immediately after spawning due to release of eggs. Thus, the change in PCB concentration due to release of eggs did not explain the higher PCB concentrations observed in males. Bioenergetics modeling results indicated that the growth dilution effect could account for males being 19% higher in PCB concentration compared with females. Thus, the bulk of the observed difference in PCB concentrations between the sexes was not explained by growth dilution. We concluded that a higher rate of energy expenditure in males, stemming from greater activity and a greater resting metabolic rate, was most likely the primary driver for the observed difference in PCB concentrations between the sexes.

Sex differences in contaminant concentrations of fish: a synthesis.

A comparison of whole-fish polychlorinated biphenyl (PCB) and total mercury (Hg) concentrations in mature males with those in mature females may provide insights into sex differences in behavior, metabolism, and other physiological processes. In eight species of fish, we observed that males exceeded females in whole-fish PCB concentration by 17 to 43 %. Based on results from hypothesis testing, we concluded that these sex differences were most likely primarily driven by a higher rate of energy expenditure, stemming from higher resting metabolic rate (or standard metabolic rate (SMR)) and higher swimming activity, in males compared with females. A higher rate of energy expenditure led to a higher rate of food consumption, which, in turn, resulted in a higher rate of PCB accumulation. For two fish species, the growth dilution effect also made a substantial contribution to the sex difference in PCB concentrations, although the higher energy expenditure rate for males was still the primary driver. Hg concentration data were available for five of the eight species. For four of these five species, the ratio of PCB concentration in males to PCB concentration in females was substantially greater than the ratio of Hg concentration in males to Hg concentration in females. In sea lamprey (Petromyzon marinus), a very primitive fish, the two ratios were nearly identical. The most plausible explanation for this pattern was that certain androgens, such as testosterone and 11-ketotestosterone, enhanced Hg-elimination rate in males. In contrast, long-term elimination of PCBs is negligible for both sexes. According to this explanation, males not only ingest Hg at a higher rate than females but also eliminate Hg at a higher rate than females, in fish species other than sea lamprey. Male sea lamprey do not possess either of the above-specified androgens. These apparent sex differences in SMRs, activities, and Hg-elimination rates in teleost fishes may also apply, to some degree, to higher vertebrates including humans. Our synthesis findings will be useful in (1) developing sex-specific bioenergetics models for fish, (2) developing sex-specific risk assessment models for exposure of humans and wildlife to contaminants, and (3) refining Hg mass balance models for fish and higher vertebrates.

Individual-Based Model for Growth of Young-of-the-Year Walleye: A Piece of the Recruitment Puzzle.

Young-of-the-year (YOY) walleye (Stizostedion vitreum vitreum) growth is a vital step in walleye recruitment. An individual-based model (IBM) was developed to describe the growth of YOY walleye in Oneida Lake (New York, USA) and Lake Mendota (Wisconsin, USA). In Oneida Lake the only prey species included in the model was yellow perch (Perca flavescens), whereas both yellow perch and bluegill (Lepomis macrochirus) were prey species in the model for Lake Mendota. IBM predictions for length frequencies of the YOY walleye population at the end of the growing season showed good agreement with observed length frequencies. A theoretical relationship was derived between the encounter rate, @l, used in the IBM and the half-saturation constant, k, used in a type II functional response model. Estimates of k from the two models showed good agreement, thus corroborating the value of ° chosen for the IBM application to Oneida Lake. The mean length of the YOY walleye cohort and the percentage of larger (@> 175 mm in total length) walleyes in the cohort at the end of the growing season were most sensitive to gross growth efficiency, bioenergetics parameters for maximum daily consumption by walleyes, and the ratio of prey length to predator length at which the prey is susceptible to predation. In Lake Mendota the vulnerability of bluegills to predation by YOY walleyes was especially important in determining the growth of walleyes during their first growing season. The IBM approach was valuable for modeling those stages of life history in which characteristics of the individual were critical in determining recruitment.

Stocking Strategies for Fingerling Walleyes: An Individual-Based Model Approach.

The success of any program for stocking walleye (Stizostedion vitreum vitreum) fingerlings is strongly dependent on growth of the stocked fish during the summer and early fall months immediately following their release into lakes, reservoirs, or rivers. An individual-based model (IBM) was developed to describe growth of young-of-the-year (YOY) walleyes in Lake Mendota (Wisconsin, USA). The IBM was used to evaluate stocking strategies for walleye fingerlings. According to the rules of this simulation model, predation by a walleye would occur only if the walleye was sufficiently large relative to the prey individual. The length-frequency distribution of the YOY walleye population at the end of the 1989 growing season was predicted accurately by the model. During 1989, walleye fingerlings with a mean total length of 50 mm were stocked into Lake Mendota on 28 June. Simulations were performed to investigate the effects of the size of stocked fingerlings and the timing of stocking on subsequent YOY walleye growth. These simulations revealed that if walleye fingerlings were stocked on 28 June, at an average total length of 60 mm rather than 50 mm, then the proportion of large (total length of @>175 mm) fish in the YOY walleye population at the end of the growing season would have increased threefold over the observed proportion. Economic cost per large walleye was minimized when average total length at stocking was 62 mm. Stocking 50-mm walleye fingerlings on 14 June instead of 28 June resulted in a tripling of the percentage of large walleyes at the end of the growing season.

Laboratory estimation of net trophic transfer efficiencies of PCB Congeners to lake trout (Salvelinus namaycush) from its prey.

A technique for laboratory estimation of net trophic transfer efficiency (γ) of polychlorinated biphenyl (PCB) congeners to piscivorous fish from their prey is described herein. During a 135-day laboratory experiment, we fed bloater (Coregonus hoyi) that had been caught in Lake Michigan to lake trout (Salvelinus namaycush) kept in eight laboratory tanks. Bloater is a natural prey for lake trout. In four of the tanks, a relatively high flow rate was used to ensure relatively high activity by the lake trout, whereas a low flow rate was used in the other four tanks, allowing for low lake trout activity. On a tank-by-tank basis, the amount of food eaten by the lake trout on each day of the experiment was recorded. Each lake trout was weighed at the start and end of the experiment. Four to nine lake trout from each of the eight tanks were sacrificed at the start of the experiment, and all 10 lake trout remaining in each of the tanks were euthanized at the end of the experiment. We determined concentrations of 75 PCB congeners in the lake trout at the start of the experiment, in the lake trout at the end of the experiment, and in bloaters fed to the lake trout during the experiment. Based on these measurements, γ was calculated for each of 75 PCB congeners in each of the eight tanks. Mean γ was calculated for each of the 75 PCB congeners for both active and inactive lake trout. Because the experiment was replicated in eight tanks, the standard error about mean γ could be estimated. Results from this type of experiment are useful in risk assessment models to predict future risk to humans and wildlife eating contaminated fish under various scenarios of environmental contamination.

Polychlorinated biphenyl congener distributions in burbot: evidence for a latitude effect.

The authors compared the distributions of the congeners of polychlorinated biphenyls (PCBs) detected in whole-body samples of burbot (Lota lota) from Great Slave Lake and Lake Erie. Total PCB concentrations in Great Slave Lake burbot were about one-sixtieth of the concentrations in Lake Erie burbot. Burbot from Great Slave Lake contained a higher proportion of lower-chlorinated (2-6 chlorines) congeners than did burbot from Lake Erie; the reverse occurred for more highly chlorinated (7-9 chlorines) congeners. Hexachloro congeners, followed by pentachloro congeners, dominated the proportions of total PCBs in burbot from both lakes. There were no differences between sexes in whole-body samples or between gonad and somatic tissues in the proportions of the 39 congeners and 3 sets of coeluters detected in burbot from Great Slave Lake. In contrast, there were distinct sex differences in congener distributions for older burbot from Lake Erie. The results generally supported a prediction of higher proportions of lower-chlorinated PCB homologs in organisms in remote polar areas. However, the latitudinal effect on PCB congener distribution may be more complex than that portrayed in previous studies.