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Steven A. Pothoven

Publications and source records attributed to Steven A. Pothoven.

31 records · Page 2Linked to original sources

Changes in the Lake Michigan food web following dreissenid mussel invasions: A synthesis

Using various available time series for Lake Michigan, we examined changes in the Lake Michigan food web following the dreissenid mussel invasions and identified those changes most likely attributable to these invasions, thereby providing a synthesis. Expansion of the quagga mussel ( Dreissena rostriformis bugensis ) population into deeper waters, which began around 2004, appeared to have a substantial predatory effect on both phytoplankton abundance and primary production, with annual primary production in offshore (> 50 m deep) waters being reduced by about 35% by 2007. Primary production likely decreased in nearshore waters as well, primarily due to predatory effects exerted by the quagga mussel expansion. The drastic decline in Diporeia abundance in Lake Michigan during the 1990s and 2000s has been attributed to dreissenid mussel effects, but the exact mechanism by which the mussels were negatively affecting Diporeia abundance remains unknown. In turn, decreased Diporeia abundance was associated with reduced condition, growth, and/or energy density in alewife ( Alosa pseudoharengus ), lake whitefish ( Coregonus clupeaformis ), deepwater sculpin ( Myoxocephalus thompsonii ), and bloater ( Coregonus hoyi ). However, lake-wide biomass of salmonines, top predators in the food web, remained high during the 2000s, and consumption of alewives by salmonines actually increased between the 1980–1995 and 1996–2011 time periods. Moreover, abundance of the lake whitefish population, which supports Lake Michigan's most valuable commercial fishery, remained at historically high levels during the 2000s. Apparently, counterbalancing mechanisms operating within the complex Lake Michigan food web have enabled salmonines and lake whitefish to retain relatively high abundances despite reduced primary production.

Lake Michigan

Coupling age-structured stock assessment and fish bioenergetics models: a system of time-varying models for quantifying piscivory patterns during the rapid trophic shift in the main basin of Lake Huron

We quantified piscivory patterns in the main basin of Lake Huron during 1984–2010 and found that the biomass transfer from prey fish to piscivores remained consistently high despite the rapid major trophic shift in the food webs. We coupled age-structured stock assessment models and fish bioenergetics models for lake trout ( Salvelinus namaycush ), Chinook salmon ( Oncorhynchus tshawytscha ), walleye ( Sander vitreus ), and lake whitefish ( Coregonus clupeaformis ). The model system also included time-varying parameters or variables of growth, length–mass relations, maturity schedules, energy density, and diets. These time-varying models reflected the dynamic connections that a fish cohort responded to year-to-year ecosystem changes at different ages and body sizes. We found that the ratio of annual predation by lake trout, Chinook salmon, and walleye combined with the biomass indices of age-1 and older alewives ( Alosa pseudoharengus ) and rainbow smelt ( Osmerus mordax ) increased more than tenfold during 1987–2010, and such increases in predation pressure were structured by relatively stable biomass of the three piscivores and stepwise declines in the biomass of alewives and rainbow smelt. The piscivore stability was supported by the use of alternative energy pathways and changes in relative composition of the three piscivores. In addition, lake whitefish became a new piscivore by feeding on round goby ( Neogobius melanostomus ). Their total fish consumption rivaled that of the other piscivores combined, although fish were still a modest proportion of their diet. Overall, the use of alternative energy pathways by piscivores allowed the increases in predation pressure on dominant diet species.

Lake Huron

Increased piscivory by lake whitefish in Lake Huron

We evaluated the diet of Lake Whitefish Coregonus clupeaformis in Lake Huron during 2002–2011 to determine the importance of Round Goby Neogobius melanostomus and other fish as prey items. Lake Whitefish that had reached approximately 400 mm in length incorporated fish into their diets. The overall percentage of adult Lake Whitefish in Lake Huron that had eaten fish increased from 10% in 2002–2006 to 20% in 2007–2011, with a corresponding decrease in the frequency of Lake Whitefish that ate Dreissena spp. from 52% to 33%. During 2002–2006, Round Goby (wet mass, 38%), sculpins (Cottidae) (34%), and Ninespine Stickleback Pungitius pungitius (18%) were the primary fish eaten, whereas Round Goby accounted for 92% of the fish eaten in 2007–2011. Overall, Round Goby were found in the fewest Lake Whitefish stomachs in the north region of Lake Huron (6%) and in the most in the central (23%) and south (19%) regions of the lake. In the central region, Round Goby were eaten during all seasons that were sampled (spring through fall). In the south region, Round Goby were eaten only in the winter and spring but not in the summer when Dreissena spp. and spiny water flea Bythotrephes longimanus dominated the diet. Based on the 2007–2011 diet composition, an individual Lake Whitefish would need to have increased their consumption relative to that in 1983–1994 by 6% in the north region, 12% in the central region, and 41% in the southern region in order to achieve the same growth that was observed before dreissenid mussels arrived. However, Lake Whitefish weight adjusted for length only increased by 2% between 2002–2006 and 2007–2011 in the central region, decreased by 4% in the northern region, and remained constant in the southern region. This suggests that a shift toward more frequent piscivory does not necessarily improve the condition of a generalist feeder like Lake Whitefish.

North American Journal of Fisheries Management

Effects of activity and energy budget balancing algorithm on laboratory performance of a fish bioenergetics model

We evaluated the performance of the Wisconsin bioenergetics model for lake trout Salvelinus namaycush that were fed ad libitum in laboratory tanks under regimes of low activity and high activity. In addition, we compared model performance under two different model algorithms: (1) balancing the lake trout energy budget on day t based on lake trout energy density on day t and (2) balancing the lake trout energy budget on day t based on lake trout energy density on day t + 1. Results indicated that the model significantly underestimated consumption for both inactive and active lake trout when algorithm 1 was used and that the degree of underestimation was similar for the two activity levels. In contrast, model performance substantially improved when using algorithm 2, as no detectable bias was found in model predictions of consumption for inactive fish and only a slight degree of overestimation was detected for active fish. The energy budget was accurately balanced by using algorithm 2 but not by using algorithm 1. Based on the results of this study, we recommend the use of algorithm 2 to estimate food consumption by fish in the field. Our study results highlight the importance of accurately accounting for changes in fish energy density when balancing the energy budget; furthermore, these results have implications for the science of evaluating fish bioenergetics model performance and for more accurate estimation of food consumption by fish in the field when fish energy density undergoes relatively rapid changes.

Transactions of the American Fisheries Society

Energy density of bloaters in the upper Great Lakes

We evaluated the energy density of bloaters Coregonus hoyi as a function of fish size across Lakes Michigan, Huron, and Superior in 2008–2009 and assessed how differences in energy density are related to factors such as biomass density of bloaters and availability of prey. Additional objectives were to compare energy density between sexes and to compare energy densities of bloaters in Lake Michigan between two time periods (1998–2001 and 2008–2009). For the cross-lake comparisons in 2008, energy density increased with fish total length (TL) only in Lake Michigan. Mean energy density adjusted for fish size was 8% higher in bloaters from Lake Superior than in bloaters from Lake Huron. Relative to fish in these two lakes, small (<125 mm TL) bloaters from Lake Michigan had lower energy density, whereas large (>175 mm TL) bloaters had higher energy density. In 2009, energy density increased with bloater size, and mean energy density adjusted for fish size was about 9% higher in Lake Michigan than in Lake Huron (Lake Superior was not sampled during 2009). Energy density of bloaters in Lake Huron was generally the lowest among lakes, reflecting the relatively low densities of opossum shrimp Mysis diluviana and the relatively high biomass of bloaters reported for that lake. Other factors, such as energy content of prey, growing season, or ontogenetic differences in energy use strategies, may also influence cross-lake variation in energy density. Mean energy density adjusted for length was 7% higher for female bloaters than for male bloaters in Lakes Michigan and Huron. In Lake Superior, energy density did not differ between males and females. Finally, energy density of bloaters in Lake Michigan was similar between the periods 2008–2009 and 1998–2001, possibly due to a low population abundance of bloaters, which could offset food availability changes linked to the loss of prey such as the amphipods Diporeia spp.

Lake Michigan;Lake Huron;Lake Superior

Influences on Bythotrephes longimanus life-history characteristics in the Great Lakes

We compared Bythotrephes population demographics and dynamics to predator (planktivorous fish) and prey (small-bodied crustacean zooplankton) densities at a site sampled through the growing season in Lakes Michigan, Huron, and Erie. Although seasonal average densities of Bythotrephes were similar across lakes (222/m 2 Erie, 247/m 2 Huron, 162/m 2 Michigan), temporal trends in abundance differed among lakes. In central Lake Erie where Bythotrephes ' prey assemblage was dominated by small individuals (60%), where planktivorous fish densities were high (14,317/ha), and where a shallow water column limited availability of a deepwater refuge, the Bythotrephes population was characterized by a small mean body size, large broods with small neonates, allocation of length increases mainly to the spine rather than to the body, and a late summer population decline. By contrast, in Lake Michigan where Bythotrephes ' prey assemblage was dominated by large individuals (72%) and planktivorous fish densities were lower (5052/ha), the Bythotrephes population was characterized by a large mean body size (i.e., 37–55% higher than in Erie), small broods with large neonates, nearly all growth in body length occurring between instars 1 and 2, and population persistence into fall. Life-history characteristics in Lake Huron tended to be intermediate to those found in Lakes Michigan and Erie, reflecting lower overall prey and predator densities (1224/ha) relative to the other lakes. Because plasticity in life history can affect interactions with other species, our findings point to the need to understand life-history variation among Great Lakes populations to improve our ability to model the dynamics of these ecosystems.

Great Lakes

Seasonal zooplankton dynamics in Lake Michigan: disentangling impacts of resource limitation, ecosystem engineering, and predation during a critical ecosystem transition

We examined seasonal dynamics of zooplankton at an offshore station in Lake Michigan from 1994 to 2003 and 2007 to 2008. This period saw variable weather, declines in planktivorous fish abundance, the introduction and expansion of dreissenid mussels, and a slow decline in total phosphorus concentrations. After the major expansion of mussels into deep water (2007–2008), chlorophyll in spring declined sharply, Secchi depth increased markedly in all seasons, and planktivorous fish biomass declined to record-low levels. Overlaying these dramatic ecosystem-level changes, the zooplankton community exhibited complex seasonal dynamics between 1994–2003 and 2007–2008. Phenology of the zooplankton maximum was affected by onset of thermal stratification, but there was no other discernable effect due to temperature. Interannual variability in zooplankton biomass during 1994 and 2003 was strongly driven by planktivorous fish abundance, particularly age-0 and age-1 alewives. In 2007–2008, there were large decreases in Diacyclops thomasi and Daphnia mendotae possibly caused by food limitation as well as increased predation and indirect negative effects from increases in Bythotrephes longimanus abundance and in foraging efficiency associated with increased light penetration. The Bythotrephes increase was likely driven in part by decreased predation from yearling and older alewife. While there was a major decrease in epilimnetic–metalimnetic herbivorous cladocerans in 2007–2008, there was an increase in large omnivorous and predacious calanoid copepods, especially those in the hypolimnion. Thus, changes to the zooplankton community are the result of cascading, synergistic interactions, including a shift from vertebrate to invertebrate planktivory and mussel ecosystem impacts on light climate and chlorophyll.

Journal of Great Lakes Research

Comparing seasonal dynamics of the Lake Huron zooplankton community between 1983-1984 and 2007 and revisiting the impact of Bythotrephes planktivory

Zooplankton community composition can be influenced by lake productivity as well as planktivory by fish or invertebrates. Previous analyses based on long-term Lake Huron zooplankton data from August reported a shift in community composition between the 1980s and 2000s: proportional biomass of calanoid copepods increased while that of cyclopoid copepods and herbivorous cladocerans decreased. Herein, we used seasonally collected data from Lake Huron in 1983–1984 and 2007 and reported similar shifts in proportional biomass. We also used a series of generalized additive models to explore differences in seasonal abundance by species and found that all three cyclopoid copepod species ( Diacyclops thomasi, Mesocylops edax, Tropocyclops prasinus mexicanus ) exhibited higher abundance in 1983–1984 than in 2007. Surprisingly, only one ( Epischura lacustris ) of seven calanoid species exhibited higher abundance in 2007. The results for cladocerans were also mixed with Bosmina spp. exhibiting higher abundance in 1983–1984, while Daphnia galeata mendotae reached a higher level of abundance in 2007. We used a subset of the 2007 data to estimate not only the vertical distribution of Bythotrephes longimanus and their prey, but also the consumption by Bythotrephes in the top 20 m of water. This epilimnetic layer was dominated by copepod copepodites and nauplii, and consumption either exceeded (Hammond Bay site) or equaled 65% (Detour site) of epilimnetic zooplankton production. The lack of spatial overlap between Bythotrephes and herbivorous cladocerans and cyclopoid copepod prey casts doubt on the hypothesis that Bythotrephes planktivory was the primary driver underlying the community composition changes in the 2000s.

Lake Huron

Indirect consequences of hypolimnetic hypoxia on zooplankton growth in a large eutrophic lake

Diel vertical migration (DVM) of some zooplankters in eutrophic lakes is often compressed during peak hypoxia. To better understand the indirect consequences of seasonal hypolimnetic hypoxia, we integrated laboratory-based experimental and field-based observational approaches to quantify how compressed DVM can affect growth of a cladoceran, Daphnia mendotae , in central Lake Erie, North America. To evaluate hypoxia tolerance of D. mendotae , we conducted a survivorship experiment with varying dissolved oxygen concentrations, which demonstrated high sensitivity of D. mendotae to hypoxia (≤2 mg O2 l −1 ), supporting the field observations of their behavioral avoidance of the hypoxic hypolimnion. To investigate the effect of temporary changes in habitat conditions associated with the compressed DVM, we quantified the growth of D. mendotae , using a 3 (food quantity) × 2 (temperature) factorial design laboratory experiment. Neither food quantity nor temperature affected short-term growth in body length of D. mendotae . However, D. mendotae RNA content (an index of short-term condition) decreased under starvation, indicating an immediate response of short-term feeding on condition. We further evaluated the effect of hypoxia-induced upward shifts in vertical distribution by quantifying the RNA content of D. mendotae from central Lake Erie before and during peak hypoxia. Despite high temperature and food quantity in the upper water column, RNA content in field-collected D. mendotae remained low during peak hypoxia. Furthermore, D. mendotae collected during peak hypoxia consisted of only small-bodied (<~1.25 mm) individuals, suggesting that behavioral avoidance of the hypoxic hypolimnion may also have indirect fitness costs.

Aquatic Biology

Effects of hypoxia on consumption, growth, and RNA:DNA ratios of young Yellow Perch

As in various freshwater and coastal marine ecosystems worldwide, seasonal bottom water hypoxia is a recurring phenomenon in Lake Erie’s central basin. While bottom hypoxia can strongly affect sessile benthic animals, its effects on mobile organisms such as fish are less understood. We evaluated the potential for bottom hypoxia to affect the growth rates of yellow perch Perca flavescens, a species of ecological and economic importance in the lake. To this end, we (1) conducted laboratory experiments to quantify the effects of reduced dissolved oxygen on consumption, somatic growth, and RNA:DNA ratios (an index of short-term growth) of young yellow perch and (2) explored the effects of bottom hypoxia on young yellow perch growth in Lake Erie’s central basin by collecting individuals in hypoxic and normoxic regions of the lake and quantifying their RNA:DNA ratios. Yellow perch consumption and growth in our experiments declined under hypoxic conditions (≤2 mg O2/L). While yellow perch RNA:DNA ratios responded strongly to experimental temperature, nucleic acid ratios were not significantly affected by dissolved oxygen or feeding ration. We did, however, observe a positive correlation between yellow perch growth and RNA:DNA ratios at low temperatures (11◦C). The nucleic acid ratios of yellow perch collected in Lake Erie varied spatiotemporally, but their patterns were not consistent with hypoxia. In short, while yellow perch consumption and growth rates respond directly and negatively to low oxygen conditions, these responses are not necessarily reflected in RNA:DNA ratios. Moreover, in central Lake Erie, where yellow perch can behaviorally avoid hypoxic areas, the RNA:DNA ratios of yellow perch do not respond strongly to bottom hypoxia. Thus, this study suggests that there is no strong negative effect of bottom hypoxia on the growth of young yellow perch in Lake Erie.

Transactions of the American Fisheries Society

Dreissenid mussels are not a "dead end" in Great Lakes food webs

Dreissenid mussels have been regarded as a “dead end” in Great Lakes food webs because the degree of predation on dreissenid mussels, on a lakewide basis, is believed to be low. Waterfowl predation on dreissenid mussels in the Great Lakes has primarily been confined to bays, and therefore its effects on the dreissenid mussel population have been localized rather than operating on a lakewide level. Based on results from a previous study, annual consumption of dreissenid mussels by the round goby (Neogobius melanostomus) population in central Lake Erie averaged only 6 kilotonnes (kt; 1 kt = one thousand metric tons) during 1995–2002. In contrast, our coupling of lake whitefish (Coregonus clupeaformis) population models with a lake whitefish bioenergetics model revealed that lake whitefish populations in Lakes Michigan and Huron consumed 109 and 820 kt, respectively, of dreissenid mussels each year. Our results indicated that lake whitefish can be an important predator on dreissenid mussels in the Great Lakes, and that dreissenid mussels do not represent a “dead end” in Great Lakes food webs. The Lake Michigan dreissenid mussel population has been estimated to be growing more than three times faster than the Lake Huron dreissenid mussel population during the 2000s. One plausible explanation for the higher population growth rate in Lake Michigan would be the substantially higher predation rate by lake whitefish on dreissenid mussels in Lake Huron.

Journal of Great Lakes Research

Changes in seasonal energy dynamics of alewife ( Alosa pseudoharengus ) in Lake Michigan after invasion of dreissenid mussels

The dreissenid mussel invasion of Lake Michigan during the 1990s has been linked to a concomitant decrease in the abundance of the amphipod Diporeia . We tracked the seasonal energy dynamics of alewife ( Alosa pseudoharengus ) in Lake Michigan during 2002&ndash;2004 and compared our findings with previously published results for years 1979&ndash;1981. Adult alewife energy density exhibited a pronounced seasonal cycle during both the pre-invasion and post-invasion periods, with energy density in October or November nearly twice as high as that in early summer. However, on average, adult alewife energy density was 23% lower during the post-invasion period compared with the pre-invasion period. This significant decline in energy density was attributable to decreased importance of Diporeia in adult alewife diet. In contrast, energy density of juvenile alewives did not significantly differ between the pre-invasion and post-invasion periods. To attain a weight of 8 kg by age 4, bioenergetics modeling indicated that a Chinook salmon ( Oncorhynchus tshawytscha ) in Lake Michigan would have to consume 22.1% more alewives during the post-invasion period compared with the pre-invasion period.

Canadian Journal of Fisheries and Aquatic Sciences

Evaluation of a lake whitefish bioenergetics model

We evaluated the Wisconsin bioenergetics model for lake whitefish Coregonus clupeaformis in the laboratory and in the field. For the laboratory evaluation, lake whitefish were fed rainbow smelt Osmerus mordax in four laboratory tanks during a 133-d experiment. Based on a comparison of bioenergetics model predictions of lake whitefish food consumption and growth with observed consumption and growth, we concluded that the bioenergetics model furnished significantly biased estimates of both food consumption and growth. On average, the model overestimated consumption by 61% and underestimated growth by 16%. The source of the bias was probably an overestimation of the respiration rate. We therefore adjusted the respiration component of the bioenergetics model to obtain a good fit of the model to the observed consumption and growth in our laboratory tanks. Based on the adjusted model, predictions of food consumption over the 133-d period fell within 5% of observed consumption in three of the four tanks and within 9% of observed consumption in the remaining tank. We used polychlorinated biphenyls (PCBs) as a tracer to evaluate model performance in the field. Based on our laboratory experiment, the efficiency with which lake whitefish retained PCBs from their food (I?) was estimated at 0.45. We applied the bioenergetics model to Lake Michigan lake whitefish and then used PCB determinations of both lake whitefish and their prey from Lake Michigan to estimate p in the field. Application of the original model to Lake Michigan lake whitefish yielded a field estimate of 0.28, implying that the original formulation of the model overestimated consumption in Lake Michigan by 61%. Application of the bioenergetics model with the adjusted respiration component resulted in a field I? estimate of 0.56, implying that this revised model underestimated consumption by 20%.

Transactions of the American Fisheries Society