Spatial and temporal trends in invertebrate communities of Great Lakes coastal wetlands,with emphasis on Saginaw Bay of Lake Huron

The shallow-sloping coastal bathymetry of Saginaw Bay (Lake Huron) supports broad fringing wetlands. Because benthic invertebrates form an important forage base for fish, wading birds, and waterfowl that utilize these habitats, understanding the drivers of invertebrate community structure has significant management implications. We used Great Lakes basin-wide data from 2002 to place Saginaw Bay wetland invertebrate communities and their environmental drivers into a basin-wide context. Various aspects of community structure were highly correlated with fetch and watershed agriculture across the basin. Saginaw Bay wetlands had relatively high fetch and watershed agriculture and supported unique invertebrate communities, typified by high abundances of many insect taxa. Wetlands from other regions around the basin tended to have more crustaceans and gastropods than the Saginaw Bay wetlands. A 1997–2012 time series from three representative Saginaw Bay wetlands revealed substantial shifts in community structure throughout the period, especially from 2001 through 2004. These years followed a 1-m decline in Lake Huron water levels that occurred between 1997 and 2000. Major community changes included decreasing insect abundance, especially chironomids, and increasing crustacean abundances, especially Hyalella azteca (Amphipoda). While factors in addition to water levels were likely also important, our time series analysis reveals the marked temporal dynamics of Saginaw Bay wetland invertebrate communities and suggests that water level decline may have influenced these communities substantially. Both the spatial and temporal community patterns that we found should be considered in future bio-assessments utilizing wetland invertebrates.     

Long-term Ecological Consequences of Anthropogenic Disturbance on Saginaw Bay Coastal Wet Meadow Vegetation

Natural disturbance related to fluctuating water levels governs the structure and composition of Saginaw Bay coastal wet meadow vegetation. European settlement introduced anthropogenic disturbance to the Saginaw Bay coastal zone, but the impact of anthropogenic disturbance on the vegetation is not known. We compared vegetation and abiotic data from six reference and six disturbed sites to examine anthropogenic impacts on Saginaw Bay coastal wet meadow vegetation. Reference sites were the least disturbed wet meadows available, whereas disturbed sites had suffered various anthropogenic impacts 10–30 years before the study started. We observed no significant difference in mean aboveground plot biomass, stem density, litter depth, hummock height and elevation, or mean site species richness, Shannon-Wiener diversity, coefficient of conservatism, and floristic quality between reference and disturbed sites. The importance values of only two of 15 major species differed significantly between reference and disturbed sites. Native wet meadow species re-vegetated previously disturbed sites, suggesting that Saginaw Bay coastal wet meadow vegetation is resilient once released from at least some forms of anthropogenic disturbance. Purple loosestrife (Lythrum salicaria L.) and several agricultural weeds occurred only in disturbed sites, suggesting a potentially useful metric for monitoring the health of Great Lakes coastal wet meadow vegetation. Additional studies are needed to gain insight into the probable revegetation trajectories of recently disturbed coastal wet meadows and to determine the form and utility of a “weedy plants” metric for classifying the condition of these wetlands.     

A status assessment and review of the herpetofauna within the Saginaw Bay of Lake Huron

Amphibians and reptiles are key bioindicators of environmental health and habitat quality and can be used to provide baseline information to help assess habitat conditions and evaluate restoration success. In 2011 and 2012 we conducted comprehensive herpetological surveys throughout the Saginaw Bay area and assessed community composition, species richness, and spatial distribution. We also compared current distributions to historic observations and habitat conditions. A total of 25 taxa (13 species of reptiles and 12 species of amphibians) were observed within the Saginaw Bay area during this study. Herpetofauna were conspicuously unobserved in areas where Phragmites australis dominates the vegetation community and were concentrated in remaining areas of suitable habitat adjacent to Phragmites. Herpetofauna observations were clustered in areas where Phragmites and other invasive plant species were rare or absent. We were able to relate categorization of Phragmites invasion (i.e., 1 = 0–10% colonization detected, 2 = 10–50%, 3 > 50%) to both a biologically and statistically significant decrease in amphibian and reptile species richness. Our results indicate that Saginaw Bay can support a diverse herpetofauna community and there is potential to restore and improve this region for rare and common amphibian and reptile species. Removal of invasive plant species would greatly improve herpetofaunal communities within Saginaw Bay. Our results will help this region's resource professionals assess the quality of habitat and set goals for restoration of amphibian and reptile habitats.     

A 200 year chronology of burrowing mayflies (Hexagenia spp.) in Saginaw Bay

After an absence of 50 years, burrowing mayflies (Hexagenia spp.) colonized western Lake Erie which led to interest in whether this fauna can be used to measure recovery in nearshore waters throughout the Great Lakes. However, in many areas we do not know if mayflies were native/endemic and thus, whether recovery is a logical measure to assess progress of recovery. In the present study, we construct a chronologic record of relative abundance of burrowing mayflies in Saginaw Bay by the use of mayfly tusks and radionuclides in sediments (i.e., a paleoecologic record) and historic records of mayfly nymphs in the bay. These records reveal that mayflies: (1) were few before 1799, which indicates that nymphs were probably native/endemic in the bay, (2) increased between 1799 and 1807 and remained at relatively high levels between 1807 and 1965, probably in response to increased nutrient run-off from the watershed, (3) declined dramatically between 1965 and 1973, probably as a result of excessive eutrophication in the mid-1950s; and, (4) were few and highly variable between 1973 and 2001, probably as a result of low and unstable abundances of mayfly nymphs. Historic records verify that nymphs disappeared in the bay in the late-1950s to early-1960s which is in agreement with the paleoecologic record. Reoccurrence of low abundances of nymphs in the bay between 1991 and 2008 and comparison of chronologic records of nymphs in Saginaw Bay and western Lake Erie suggest that mayflies may return to Saginaw Bay in the early-21st century. Undoubtedly, watershed conservation and three decades of pollution abatement have set the stage for a recovery of burrowing mayflies in Saginaw Bay, and possibly in other areas of the Great Lakes.     

Modeling the implications of multiple hatching sites for larval dynamics in the resurgent Saginaw Bay walleye population

The early life environment experienced by most larval fish is largely dependent on a combination of hatch site and water currents. Until larvae are able to swim fast enough to overcome currents, they are largely passively transported and have limited control over ambient environmental conditions, including temperature and prey availability. These factors strongly influence growth and survival of larvae, with direct consequences for subsequent recruitment. Early life survival of Saginaw Bay walleye was formerly limited by alewife predation on larvae; but following the collapse of Lake Huron alewives, the walleye population has rebounded and recruitment success may now be influenced by other factors including spawning habitat. We sought to assess the implications of successful hatching at multiple locations in Saginaw Bay, using a hydrodynamics model, particle transport model, and an individual-based bioenergetics model in series. Model results were compared to locations of young larvae collected in Saginaw Bay during 2009–2010. Results suggest that larval growth is strongly influenced by hatch date, driven by seasonal variation in temperature between sites. Larvae hatched at any location could be transported extensively within inner Saginaw Bay before reaching a sufficient size to swim independently of currents, and retention within the productive inner bay varied among years and sites. Our results indicate multiple larval walleye origins in the field, augmenting the continued production from the Saginaw River system. Successful hatching at more locations would serve to buffer walleye recruitment variation through portfolio effects, supporting arguments for more emphasis on diverse spawning habitat management and restoration.     

Assessment of remnant unionid assemblages in a selection of Great Lakes coastal wetlands

Populations of native unionids have been in steady decline over the past century. The invasion of dreissenid mussels (Dreissena polymorpha and Dreissena rostriformis bugensis) in the mid-1980's impacted already imperiled unionid populations by greatly increasing their regional extinction rates. A selection of Great Lakes coastal wetlands around Michigan was surveyed to locate remnant populations of native unionids. Physical and chemical parameters were measured in coastal wetlands to evaluate the importance of these habitat parameters to remnant unionid assemblages. We assessed fouling rates by dreissenids on unionids and used artificial substrates to estimate dreissenid colonization densities. Live unionids were found in coastal wetlands of the Les Cheneaux Islands, the Lake St. Clair delta, and North Maumee Bay with significantly higher unionid fouling in the Les Cheneaux Islands compared to the other two sampling areas (F_2,76 = 4.97, p = 0.0095). No live unionids were documented in Beaver Island, Garden Island, Grand Traverse Bay, or Saginaw Bay wetlands. Dreissena colonization densities on artificial substrates averaged 19,213 m^− 2 at one site in North Maumee Bay, and 10,425 m^− 2 in Saginaw Bay, but no colonization occurred in the wetlands of Beaver Island, Garden Island, the Les Cheneaux Islands, or Grand Traverse Bay while Dreissena presence in the open water of these regions was evident. Dreissena colonization densities on artificial substrates increased with measures of anthropogenic disturbance and decreased with higher water level fluctuations and aerial exposure. Specific conductance, turbidity, and magnitude of water level fluctuations were important predictors of Dreissena colonization on artificial substrates.     

Measuring the economic benefits of Saginaw Bay coastal marsh with revealed and stated preference methods

We estimate the economic benefits of Saginaw Bay coastal marsh with the travel cost and contingent valuation methods. The travel cost method is based on revealed preferences: actual recreation behavior. Using a sample of the general population of Michigan and Michigan hunting and fishing license holders we find that Saginaw Bay recreation site selection is negatively related to travel cost and positively related to wetland acreage. The contingent valuation method is based on stated preferences: answers to hypothetical survey questions. We find that willingness-to-pay is negatively related to marsh protection cost and positively related to income and environmental organization membership. Using a combination of theory and empirical results we argue that revealed and stated preference methods are complementary when estimating the total value of coastal marsh. The present value of each acre of coastal marsh is $1870 for the purpose of recreation. The present value to recreation nonusers adds $551 per acre. The total present value of each acre of coastal marsh could be as high as $2421.     

Effects of Zebra Mussel (Dreissena polymorpha) Colonization on Water Quality Parameters in Saginaw Bay,Lake Huron

A large-scale study of Saginaw Bay was initiated in 1990 and continued through 1993 to examine the effects of the zebra mussel colonization which began in summer/fall 1991. Saginaw Bay responded quickly to the zebra mussel colonization, as fall 1991 values of chlorophyll were similar to 1992 and 1993 values. In inner Saginaw Bay, where most zebra mussels were found, chlorophyll, kPAR, and total phosphorus values decreased, and Secchi disk depth increased during the study period, regardless of the presence or absence of zebra mussels at a specific station. At outer bay control stations no significant differences were found for chlorophyll, kPAR, and Secchi disk values. In order to examine longer-term trends, water quality data from 1979–1980 (STORET) were combined with our 1990 data (pre-zebra mussel period) and compared to values from the post zebra mussel period (fall 1991, all 1992 and 1993). At stations with high densities of zebra mussels, chlorophyll and total P decreased by 66% and 48%, respectively, and Secchi disk values increased 88%. At outer bay control stations no significant differences were found for chlorophyll or Secchi disk. When parameters were averaged throughout inner Saginaw Bay, zebra mussels caused a 59% and 43% decrease in chlorophyll and in total phosphorus and a 60% increase in Secchi disk transparency. Although zebra mussels significantly altered water quality parameters in the pelagic region of Saginaw Bay, they did not necessarily change system trophic state; rather they altered the spatial partitioning of resources.     

The relative impacts of nutrient loads and invasive species on a Great Lakes food web: An Ecopath with Ecosim analysis

Excessive nutrient loads and species invasions pose significant threats to productivity and function of Great Lakes aquatic ecosystems. We used an Ecopath and Ecosim model to analyze impacts of changes in phosphorus loads, and dreissenid mussels and alewife biomass on the Saginaw Bay food web, Lake Huron. We configured the food web model in Ecopath with pre-dreissenid (1990) data on organism biomass, production, consumption, and diet from federal and state agency surveys and other sources. We conducted 70-year simulations in Ecosim of single factors (nutrients, alewives, and dreissenids) and their combinations. Phosphorus load scenarios were run with high (1970s), average (current), and low (target) levels; alewife scenarios were run with double the 1990–2003 average biomass, 1990–2003 average biomass, and alewife absence; dreissenid scenarios were run with the 1990–1996 average biomass, current (2009–2010) biomass, and dreissenid absence. Results indicated that phosphorus loads were positively correlated with simulated biomass of most food web groups, and alewife biomass was negatively correlated with biomass of most fish groups and macrozooplankton. Dreissenid impacts were most severe on lower trophic levels but were relatively minor for fish groups compared with nutrient and alewife scenarios. Dreissenids had little effect on fish because Chironomids, which feed on detritus and are the main component of fish diets, were not affected by dreissenids. Our results suggest that, under current conditions of absence of alewives and reduced dreissenid biomass, the target nutrient loads established in 1978 would not sustain current fishery harvests in Saginaw Bay given food web changes caused by invasive species.     

Trends in the distribution and abundance of Hexagenia spp. in Saginaw Bay,Lake Huron, 1954–2012: Moving towards recovery?

Since at least the 1940s, multiple anthropogenic disturbances to the Laurentian Great Lakes have had detrimental effects on benthic habitats and biota including decimating the environmentally sensitive burrowing mayfly genus Hexagenia around the mid-1950s. While remediation efforts have facilitated recovery of some populations, benthic surveys in Saginaw Bay, Lake Huron in the last 50 years have only occasionally discovered Hexagenia nymphs. Recently, adult Hexagenia swarms have been reported near the bay; therefore, we corroborated the local presence of Hexagenia adults and evaluated the current status of Saginaw Bay Hexagenia nymphs. We quantified adults during mayfly emergence events in 2010 at three Tawas City, Michigan, USA area locations, and found > 17 Hexagenia/m²/site. We quantified nymphs from Ponar grab samples collected at 57 sites in Saginaw Bay between 2009 and 2012, and found 1.5 nymphs/m² overall with nymphs present at 15.8% of sites sampled, their greatest documented distribution in Saginaw Bay since 1956. Additionally, we mapped bay sediment composition and related sampling site abiotic conditions with both Hexagenia presence and abundance using Zero-Inflated Poisson regression. Model results indicate that the probability of observed Hexagenia absence being true absence is positively related to both sediment sandiness and surficial dissolved oxygen concentration while Hexagenia abundance is greatest where surficial temperatures are ~ 18.6 °C and is also related to sediment sand content. The documentation of nearby adults and in bay nymphs may indicate the beginning of a Hexagenia return to Saginaw Bay, and, therefore, a possible improvement of the ecosystem's benthic health.     

Petroleum Hydrocarbons in Sediments of Saginaw Bay,Lake Huron

The history of petroleum contamination of Saginaw Bay is given by a 60 cm core recording sediment accumulation over the last two centuries. Aliphatic hydrocarbon concentrations average 385 μgm/gm in modern sediments and progressively decrease from 10 cm to 40 cm in the core to average 30 μgm/gm in sediments deposited before 1876. An unresolved complex mixture of hydrocarbons diagnostic of petroleum represents about 90 percent of the total concentration in all core sections deposited over the last 100 years. In older sediment, this decreases but still comprises 40 percent of the total at the bottom of the core. These patterns indicate a large input of nonbiogenic hydrocarbons which has accumulated in the sediments of Saginaw Bay. While a portion of these may originate from natural sources, such as forest fires and erosion of ancient sediments, we conservatively estimate that 2 to 4 X 10⁴ Tonne of petroleum hydrocarbons have been introduced to Saginaw Bay since the 1870s.     

Use of wetland versus open habitats by round gobies in lakes Michigan and Huron: Patterns of CPUE,length, and maturity

It has been suggested that some Great Lakes coastal wetlands may be resistant to invasion by several non-indigenous species including round goby, Neogobius melanostomus. However, there is inconclusive evidence regarding how susceptible exposed fringing coastal wetlands, in particular, are to round goby invasion. Therefore, we quantified round goby catch per unit effort (CPUE) using fyke nets in the Beaver Archipelago of Lake Michigan, and the Les Cheneaux islands and Saginaw Bay regions of Lake Huron. In addition, we examined the influence of body size and maturity on round goby habitat use. Catch per unit effort from fyke nets was highest in the Beaver Archipelago, where wetlands were dominated by small, immature round gobies and open water habitats were dominated by large adults. Fyke net catches within Les Cheneaux sites were similar between habitats and differences in size and maturity were not observed. Conversely, very few round goby were captured in wetlands of Saginaw Bay where CPUE was moderate in open water. This indicates that some exposed fringing wetlands in the Great Lakes, specifically those with high productivity, could have a higher degree of resistance to round goby invasion.     

Sex difference in polybrominated diphenyl ether concentrations of walleyes

Polybrominated diphenyl ether (PBDE) concentrations were determined for mature male and mature female walleyes (Sander vitreus) sampled from the Saginaw Bay population during 2007. PBDE concentrations in prey fish caught in the Saginaw River, the primary tributary to Saginaw Bay, and in Saginaw Bay during 2005 and 2007 also were determined. Mature male and mature female walleyes averaged 70.3 ng/g and 24.8 ng/g, respectively, in ΣPBDE, which was equal to the sum of concentrations of six PBDE congeners (BDE-28, BDE-47, BDE-99, BDE-100, BDE-153, and BDE-154). This sex difference was likely due to males spending more time in the Saginaw River system than females. Prey fish captured in the Saginaw River were roughly ten times higher in ΣPBDE than those caught in Saginaw Bay. BDE-47 was the predominant congener in both walleyes and prey fish, and this congener contributed about 50%, on average, to ΣPBDE. Congener profiles differed significantly between the two sexes of walleyes. In contrast, congener profiles of the prey fish did not differ significantly between the river-caught fish and the bay-caught fish. One plausible explanation for these congener profile results was that net trophic transfer efficiencies of PBDEs to walleyes from their prey were similar for all congeners except BDE-28, and that diet composition differed between the two sexes of walleyes.     

Colonization of the Laurentian Great Lakes by the Amphipod Gammarus tigrinus,a Native of the North American Atlantic Coast

Gammarus tigrinus, whose natural distribution is restricted to the North American Atlantic coast, has been found at numerous localities across the Laurentian Great Lakes. This amphipod was first discovered in Saginaw Bay of Lake Huron in 2002. However, analysis of archived samples and new material collected during 2001–2004 revealed that G. tigrinus is present in all of the Great Lakes. During August 2002, it occurred at an average density of 283 individuals·m⁻² in Saginaw Bay, where it was outnumbered by the resident amphipods G. fasciatus and Hyalella azteca. In terms of frequency of occurrence, G. tigrinus was the second most numerous amphipod in beds of Typha in lower Great Lakes coastal wetlands during July 2004, being outnumbered only by native G. pseudolimnaeus. Gammarus tigrinus has a history of ballast water transfer in Europe and it likely exploited this transport vector during its recent colonization of the Great Lakes.     

Movement and Loadings of Inorganic Contaminants Through the Lower Saginaw River

The objective of this study was to investigate the movement of selected contaminants (four heavy metals) and conventional limnological variables through the lower Saginaw River. We related limnological variables to concentrations of contaminants, determined if sources of these contaminants existed in the lower 8 km of the river, and calculated loadings of materials to Saginaw Bay. Concentrations of most variables changed significantly among the six sampling dates in 1990–1991, while relatively few variables were different among the six sampling stations. Causes of changes in concentrations of variables in the river over time appeared to stem from two factors, seasonality and river discharge. One phase of the sampling program was conducted on six different dates which encompassed river flows ranging from greater than 700 m³ sec⁻¹ to less than 25 m³ sec⁻¹. The date on which the least river discharge was observed included a flow reversal when Saginaw Bay water intruded into the lower Saginaw River. Statistical analyses showed that temporal variation among variables was due, in part, to fluctuating river discharge and in part to seasonal factors. The relative importance of these two components varied among parameters. For example, total suspended solids were positively correlated with river discharge, while chlorophyll varied more seasonally. In contrast to temporal trends, few parameters varied significantly among the six sampling stations. This was especially true for dissolved and paniculate metals. Loadings of key materials to Saginaw Bay from the Saginaw River were generally positively correlated with river discharge. We concluded that most substances entered the river upstream of our study section in the lower 8 km of the Saginaw River.     

Phosphorus targets and eutrophication objectives in Saginaw Bay: A 35 year assessment

An aggregated view of total phosphorus and chlorophyll a in Saginaw Bay indicates that concentrations of both constituents declined approximately in concert with declining total phosphorus (P) loads stabilizing by the late 1980s. A more spatially focused view reveals that total phosphorus declines outside of the Saginaw River plume, accompanied by more subtle chlorophyll a decreases. In contrast, soluble reactive phosphorus and ammonia have recently declined throughout the bay, while nitrate has remained relatively stable. Concentration data from nearshore transects do not exhibit large differences from open-water sample sites. The 440 tonne P/year target phosphorus load established in the 1978 amendments to the Great Lakes Water Quality Agreement has almost never been met, and total phosphorus concentrations regularly exceed the 15 μg/L concentration objective proposed in documentation supporting the 1978 amendments. Seasonal patterns in both total phosphorus and chlorophyll a are more pronounced in the most recent data, with peaks occurring in September–October. This apparently evolving seasonal pattern may result from seasonal changes in Saginaw River flow inputs, or seasonal variation in dreissenid mussel feeding and filtration rates. The adaptive management framework stipulated in the 2012 Great Lakes Water Quality Protocol should promote better monitoring of Saginaw Bay water quality into the future, with enhanced opportunities to better understand the factors that have maintained ongoing eutrophication symptoms.     

Contemporary issues in watershed and water quality modeling for eutrophication control

Three issues are discussed: controllability of nonpoint nutrient loadings using watershed models; the sometimes counter intuitive results from eutrophication models from nutrient controls for coastal waters; and the potential significant interaction of improvement in habitat for suspension feeding bivalves. For the Chesapeake Bay watershed model, and for Limit of Technology (LOT) controls, a 16% and 45% reduction in nitrogen and phosphorus, respectively, is calculated. For the Bay, it is concluded that removal of phosphorus only is less effective than nitrogen in improving bottom water DO because of differential transport of nitrogen downstream. For the Delaware estuary, a significant decline in phytoplankton chlorophyll has been observed in the absence of any nutrient controls but in the presence of improved DO. A simple model is offered that hypothesized an increase in benthic bivalve filtration of overlying water as a result of improvement in DO.     

Causes of phytoplankton changes in Saginaw Bay,Lake Huron,during the zebra mussel invasion

Colonization of the Laurentian Great Lakes by the invasive mussel Dreissena polymorpha was a significant ecological disturbance. The invasion reached Saginaw Bay, Lake Huron, in 1991 and initially cleared the waters and lowered algal biomass. However, an unexpected result occurred 3 years after the initial invasion with the return of nuisance summer blooms of cyanobacteria, a problem that had been successfully addressed with the implementation of phosphorus controls in the late 1970s. A multi-class phytoplankton model was developed and tested against field observations and then used to explore the causes of these temporal changes. Model scenarios suggest that changes in the phytoplankton community can be linked to three zebra mussel-mediated effects: (1) removal of particles resulting in clearer water, (2) increased recycle of available phosphorus throughout the summer, and (3) selective rejection of certain Microcystis strains. Light inhibition of certain phytoplankton assemblages and the subsequent alteration of competitive dynamics is a novel result of this model. These results enhance our understanding of the significant role of zebra mussels in altering lower trophic level dynamics of Saginaw Bay and suggest that their physical re-engineering of the aquatic environment was the major force driving changes in the phytoplankton community composition.     

Gender difference in walleye PCB concentrations persists following remedial dredging

Eleven male walleyes (Sander vitreus) and 10 female walleyes from the Saginaw Bay (Lake Huron) population were caught during the spawning run at Dow Dam (Midland, Michigan) in the Tittabawassee River during April 1996, and individual whole-fish polychlorinated biphenyl (PCB) determinations were made. Total PCB concentrations averaged 7.95 and 3.17 mg/kg for males and females, respectively. As part of the Natural Resource Damage Assessment remediation process, contaminated sediments from the Saginaw River, the main tributary to Saginaw Bay, were removed during 2000 and 2001. Total PCB concentrations of 10 male and 10 female walleyes caught at Dow Dam during April 2007 averaged 1.58 and 0.55 mg/kg, respectively. Thus, dredging of the Saginaw River appeared to be effective in reducing PCB concentrations of Saginaw Bay adult walleyes, as both males and females decreased in PCB concentration by more than 80% between 1996 and 2007. However, the ratio of male PCB concentration to female PCB concentration did not decline between 1996 and 2007. This persistent gender difference in PCB concentrations was apparently due to a gender difference in habitat utilization coupled with a persistent spatial gradient in prey fish PCB concentrations from the Saginaw River to Lake Huron.