Steven A. Pothoven

Changes in Diet and Body Condition of Lake Whitefish in Southern Lake Michigan Associated with Changes in Benthos

Abstract We evaluated the long-term trends of the benthic macroinvertebrate community (1980–1999) and biological attributes of lake whitefish Coregonus clupeaformis (1985–1999) in southeastern Lake Michigan. We also determined what food types were important to lake whitefish in an area where the amphipod Diporeia had not yet declined in 1998 and how the diet of lake whitefish changed as Diporeia declined during 1999–2000. Zebra mussels Dreissena polymorpha invaded the study area in 1992; Diporeia began to decline in 1993 and was nearly absent by 1999. The body condition of lake whitefish decreased after 1993 and remained low thereafter. The length at age and weight at age of lake whitefish was lower in 1992–1999 than in 1985–1991. After declines of Diporeia off the city of Muskegon, Michigan, between 1998 and 1999–2000, the proportion of Diporeia in the diet by weight fell from 70% to 25% and the percent occurrence decreased from 81% to 45%. In contrast, the proportion of lake whitefish that ate other pre...

Recent changes in primary production and phytoplankton in the offshore region of southeastern Lake Michigan

ABSTRACT Phytoplankton abundance, composition, and productivity were monitored on a bi-weekly basis from March/ April through November/December at two offshore stations in southeastern Lake Michigan in 1983–1987, 1995– 1998 and 2007–2008 (exception 1983–1984 which were sampled from May to August). During the spring isothermal mixing period, surface-mixed layer (SML) chlorophyll a and phytoplankton biomass (carbon) and water column primary productivity decreased substantially in 2007–2008 as compared to 1995–1998 (66%, 87%, and 70% decrease, respectively). Smaller or no decreases were noted between 1983–1987 and 1995–1998 (chlorophyll a 23% decrease, phytoplankton biomass 5% increase, and production 22% decrease). Phytoplankton composition also changed during the spring isothermal mixing period in 2007–2008 as compared to 1983–1987 and 1995–1998; all phytoplankton groups with the exception of cyanobacteria and chlorophytes exhibited dramatic reductions in 2007–2008. The pronounced changes in phytoplankton properties during spring mixing in 2007–2008 were attributed to the filtering activities of the quagga mussel (Dreissena rostriformis bugensis). During mid- and late thermal stratification periods, SML phytoplankton chlorophyll a and phytoplankton carbon and water column primary production exhibited only one significant change across all decades (mid-stratification production in 2007–2008 as compared to 1995–1998 and 1983–1987). Phytoplankton compositional changes in the SML also were limited during thermal stratification. The size of the deep chlorophyll layer (DCL) in 2007–2008 was similar to or smaller than those in 1983–1987 and 1995–1998. However, phytoplankton composition in the DCL changed as net diatoms constituted < 5% of total phytoplankton in the 2007–2008 DCL but over 50% in 1983–1987 and 1995–1998.

Dreissena and the disappearance of the spring phytoplankton bloom in Lake Michigan

ABSTRACT We determined the clearance rates of the profunda morph of the quagga mussel (Dreissena bugensis) using seston and Cryptomonas ozolini, a high-quality algal food, for the temperature range 1–7°C, which is the full temperature range this morph is likely to experience during isothermal conditions or in the hypolimnion of deep lakes. Experiments at 3 °C with the shallow-water morph of the quagga and the zebra mussel provided very similar results. The clearance rates were combined with dreissenid abundance in 0–30 m, 30–50 m, 50–90 m, and >90 m depth zones of the southern basin of Lake Michigan to calculate a maximum (using Cryptomonas) and minimum (using seston) fraction of the water column cleared (FC) per day in the different depth zones at 3 °C to determine dreissenid impact on the spring phytoplankton bloom from 1994 to 2008. Starting in 2003 or 2004 with the replacement of zebra mussels by quagga mussels in shallow water and expansion of quagga mussel biomass in deep water, FC began to exceed likely phytoplankton growth in the 30–50 m zone. In 2007–2008, FC greatly exceeded likely phytoplankton growth by a factor of about 5 in the 30- to 50-m depth zone, where dreissenids were extremely abundant. Low FC in the offshore region led to the hypothesis of a mid-depth carbon (C) and phosphorous (P) sink caused by mussel uptake of seston-associated C and P that affected not only the mid-depth region, but also the offshore region “downstream” of the mid-depth zone.

Long-term Trends in Benthic Macroinvertebrate Populations in Lake Huron over the Past Four Decades

ABSTRACT Surveys of the benthic macroinvertebrate community were conducted in the main basin of Lake Huron in 2000 and 2003, and in Georgian Bay and North Channel in 2002. Results were compared to surveys conducted in the 1960s and early 1970s. Although data of earlier surveys were inconsistent, our best estimates suggest that total density of the four major benthic taxa (Diporeia spp., Oligochaeta, Sphaeriidae, and Chironomidae) in the main basin declined dramatically between the early 1970s and 2000. Populations of all major taxa continued to decline between 2000 and 2003, particularly Diporeia and Sphaeriidae. Diporeia was rare or absent in the southern end of the lake and in some nearshore areas in 2000, and by 2003 was not found at depths < 50 m except in the far northeastern end of the lake. Densities of the major taxa in Georgian Bay and North Channel in 2002 were not different from densities in 1973 despite differences in survey methods. A limited study in southern Georgian Bay, however, found that densities of both Diporeia and Sphaeriidae declined to zero at most sites between 2000 and 2004. The population of Dreissena polymorpha was stable in all lake areas, but Dreissena bugensis increased, particularly at the 31–50 m depth interval in the main basin. Since there were no extensive surveys in Lake Huron in the period between nutrient abatement (late 1970s) and the establishment of Dreissena (early 1990s), it is difficult to determine relative roles of these events on observed declines. However, since phosphorus loads have been stable since the early 1980s, declines between 2000 and 2003 can likely be attributed to Dreissena.

Recent changes in density, biomass, recruitment, size structure, and nutritional state of Dreissena populations in southern Lake Michigan

ABSTRACT Trends in density, biomass, population structure, and nutritional state of Dreissena polymorpha and Dreissena rostriformis bugensis were examined in southern Lake Michigan between the 1990s and 2008. Density and biomass of D. polymorpha increased to a peak in the early 2000s and then declined. In contrast, D. rostriformis bugensis was first found in the southern basin in 2001 and has continued to increase in density or biomass at all depths ever since. In 2008, maximum mean density of D. rostriformis bugensis occurred at 16–30m (19,000/m2), but maximum biomass (AFDW) occurred at 31–50 m (43.9 g/m2). D. rostriformis bugensis has only recently (since 2005) began to increase at depths >50 m. When both species were present in 2004 at depths <50 m, a condition index (CI) for D. rostriformis bugensis was 27% higher, and shell weight per shell length was 48% lower compared to D. polymorpha. For D. rostriformis bugensis, CI decreased in 2008 compared to 2004 at 25 m and 45 m, but biochemical content (lipid, glycogen) did not. Seasonal changes in both RNA/DNA ratio (growth) and ETS (metabolic activity) in D. rostriformis bugensis were unaffected by reproductive activity, and only ETS appeared to change seasonally relative to bottom temperatures. Spawning of D. rostriformis bugensis occurred in late summer at 25 m, but occurred in spring at 45 m. Veliger densities peaked in both spring and late summer at both depths. Future population expansion (biomass) is expected to be most rapid at depths >50 m.

Changes in Consumption by Alewives and Lake Whitefish after Dreissenid Mussel Invasions in Lakes Michigan and Huron

Abstract Growth of alewives Alosa pseudoharengus and lake whitefish Coregonus clupeaformis has declined since the arrival and spread of dreissenid mussels in Lakes Michigan and Huron. Alewives are the main forage for the salmonids in Lake Michigan, and lake whitefish are the most important commercial species in both lakes. Bioenergetics modeling was used to determine consumption by the average individual fish before and after the dreissenid invasion and to provide insight into the invasions effects on fish growth and food web dynamics. Alewives feed on both zooplankton and benthic macroinvertebrates, and lake whitefish are benthivores. Annual consumption of zooplankton by an average alewife in Lake Michigan was 37% lower and consumption of benthic macroinvertebrates (amphipods Diporeia spp., opossum shrimp Mysis relicta, and Chironomidae) was 19% lower during the postinvasion period (1995–2005) than during the preinvasion period (1983–1994). Reduced consumption by alewives corresponded with reduced alewi...

Influence of Diporeia Density on Diet Composition, Relative Abundance, and Energy Density of Planktivorous Fishes in Southeast Lake Michigan

Abstract The benthic amphipod Diporeia spp. is an important prey for many fish in offshore areas of the Great Lakes, but its abundance has been rapidly decreasing. To assess the influence of Diporeia availability on the food habits, relative abundance, and energetics of planktivorous fish, the diet composition, catch per unit effort (CPUE), and energy density of plantkivorous fish in southeast Lake Michigan during 2000–2001 were compared among locations with different Diporeia densities. Diporeia densities at St. Joseph, Michigan, were near 0/m2 over much of the bottom but averaged more than 3,800/m2 at Muskegon and Little Sable Point, Michigan. Consistent with these differences in Diporeia density, fish diet composition, CPUE, and energy density varied spatially. For example, alternative prey types comprised a larger fraction of the diets of bloater Coregonus hoyi, large (>100 mm total length) alewife Alosa pseudoharengus, and slimy sculpin Cottus cognatus at St. Joseph than at Muskegon and Little Sable ...

Long-Term and Recent Changes in Southern Lake Michigan Water Quality with Implications for Present Trophic Status

ABSTRACT Southern Lake Michigan has changed in response to alterations in nutrients and invasive species. NOAA and EPA monitoring results are used to examine those changes. NOAA provides detailed seasonal resolution, but limited spatial coverage, whereas the EPA provides more spatial coverage, but limited seasonal resolution. We compare changes in total phosphorus (TP), silica, nitrate plus nitrite, and chlorophyll concentrations from before and after the invasion by the quagga mussel (Dreissena rostriformis bugensis). Although TP at NOAA stations was consistently higher than at EPA stations, both confirm declines in spring and summer surface mixed layer (SML) conditions. Chlorophyll differed at EPA and NOAA stations before quagga mussel invasion, but not after the invasion. Spring chlorophyll decreased at NOAA stations after the invasion, but summer conditions did not change at either set of stations. Pre-invasion silica at NOAA stations was slightly higher than at EPA stations, and the lakes Si reservoir increased over the study period. Basin-scale spring Si increased gradually, whereas summer SML Si increased dramatically after 2003, likely reflecting reduced diatom production. Basin-scale nitrate increased significantly from pre- to post-invasion in both spring and summer. Summer nitrate utilization declined drastically in recent years, likely reflecting reduced phytoplankton production. TP loads decreased; however, the timing of changes in chlorophyll and Si and nitrate utilization suggest the recent increase in dreissenid filtering dramatically reduced spring phytoplankton abundance and production across the entire southern basin. The offshore pelagic zone of the historically mesotrophic southern Lake Michigan is now similar to oligotrophic Lake Superior.

Evaluation of a Lake Whitefish Bioenergetics Model

Abstract 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...

Introduction of the Ponto-Caspian Bloody-red Mysid Hemimysis anomala into the Lake Michigan Basin

ABSTRACT Hemimysis anomala G.O. Sars, 1907, a mysid species native to the Ponto-Caspian region, was discovered during fall 2006 in the Lake Michigan basin. Large numbers of individuals formed aggregations (averaging 1,540 ± 333 individuals/m2) in a shallow docking basin connected to the channel linking Lake Michigan and Muskegon Lake. The population included females (63%), males (35%), and juveniles (2%). The global invasion pattern in H. anomala is similar to that in another Ponto-Caspian peracarid crustacean, Echinogammarus ischnus. As with E. ischnus, the expansion of H. anomala in North America is anticipated.