Watershed and lake influences on the energetic base of coastal wetland food webs across the Great Lakes Basin

We examined factors that influence the energy base of Great Lakes coastal wetland food webs across a basin-wide gradient of landscape disturbance. Wetland nutrient concentrations were positively correlated with a principal components-based metric of agricultural practices. Hydraulic residence time influenced the energy base of wetland food webs, with high residence-time systems based mostly on plankton and low residence-time systems based mostly upon benthos. In systems with plankton, the importance of planktonic carbon to the resident fish community generally increased with residence time. A stronger relationship was apparent with an index of nutrient loading that combined residence time and nutrient concentration as the predictor (R² = 0.289, p = 0.026). Shifts toward plankton-based food webs occurred at relatively low levels of loading. In riverine wetlands without plankton, contributions of detrital carbon to fish communities decreased significantly in response to watershed disturbance that reflected nutrient loading. In a third class of wetlands the wetland-resident fish communities were not entirely supported by within-wetland carbon sources and were significantly subsidized by nearshore habitats, which provided 35 (± 22) to 73 (± 9) % of fish community carbon. When lake-run migrant fish were included in the analyses, nearshore subsidies to all 30 wetland food webs ranged from 3 (± 2) to 79 (± 12) %. We obtained similar ranges when examining nearshore contributions to a single wetland species, northern pike. These results illustrate the spatial scale and the degree to which the energetics of coastal wetland food webs are influenced by interactions with their watersheds and Great Lakes.     

Perchlorate in environmental waters of the Laurentian Great Lakes watershed: Evidence for uneven loading

A database of nearly 500 analyses of perchlorate in water samples from the Laurentian Great Lakes (LGL) watershed is presented, including samples from streams, from the Great Lakes and their connecting waters, with a special emphasis on Lake Erie. These data were assessed to test an earlier hypothesis that loading of perchlorate to the LGL watershed is relatively uniform. Higher perchlorate concentrations in streams in more developed and urban areas appear to indicate higher rates of loading from anthropogenic sources in these areas. Variable perchlorate concentrations in samples from Lake Erie indicate transient (un-mixed) conditions, and suggest loss by microbial degradation, focused in the central basin of that lake. Interpretation of the data included estimation of annual loading by streams in various sub-watersheds, and simulations (steady state and transient state) of the mass balance of perchlorate in the Great Lakes. The results suggest uneven loading from atmospheric deposition and other sources.     

Researcher disciplines and the assessment techniques used to evaluate Laurentian Great Lakes coastal ecosystems

The Laurentian Great Lakes of North America have been a focus of environmental and ecosystem research since the Great Lakes Water Quality Agreement in 1972. This study provides a review of scientific literature directed at the assessment of Laurentian Great Lakes coastal ecosystems. Our aim was to understand the methods employed to quantify disturbance and ecosystem quality within Laurentian Great Lakes coastal ecosystems within the last 20 years. We focused specifically on evidence of multidisciplinary articles, in authorship or types of assessment parameters used. We sought to uncover: 1) where Laurentian Great Lakes coastal ecosystems are investigated, 2) how patterns in the disciplines of researchers have shifted over time, 3) how measured parameters differed among disciplines, and 4) which parameters were used most often. Results indicate research was conducted almost evenly across the five Laurentian Great Lakes and that publication of coastal ecosystems studies increased dramatically ten years after the first State of the Great Lakes Ecosystem Conference in 1994. Research authored by environmental scientists and by multiple disciplines (multidisciplinary) have become more prevalent since 2003. This study supports the likelihood that communication and knowledge-sharing is happening between disciplines on some level. Multidisciplinary or environmental science articles were the most inclusive of parameters from different disciplines, but every discipline seemed to include chemical parameters less often than biota, physical, and spatial parameters. There is a need for an increased understanding of minor nutrient, toxin, and heavy metal impacts and use of spatial metrics in Laurentian Great Lakes coastal ecosystems.     

Fish movement and migration studies in the Laurentian Great Lakes: Research trends and knowledge gaps

Resource management agencies in the Laurentian Great Lakes routinely conduct studies of fish movement and migration to understand the temporal and spatial distribution of fishes within and between the lakes and their tributaries. This literature has never been summarized and evaluated to identify common themes and future research opportunities. We reviewed 112 studies, published between 1952 and 2010, with the goal of summarizing existing research on the movement and migration of fishes in the Laurentian Great Lakes. The most commonly studied species were Lake Trout (Salvelinus namaycush), Walleye (Sander vitreus), and Lake Sturgeon (Acipenser fulvescens). Studies relied mainly on mark-recapture techniques with comparatively few using newer technologies such as biotelemetry, hydroacoustics, or otolith microchemistry/isotope analysis. Most movement studies addressed questions related to reproductive biology, effects of environmental factors on movement, stocking, and habitat use. Movement-related knowledge gaps were identified through the literature synthesis and a survey distributed to Great Lakes fisheries managers. Future studies on emigration/immigration of fishes through lake corridors, the dispersal of stocked fishes and of stock mixing were identified as being particularly important given their potential for developing lake- or region-wide harvest regulations and stocking strategies. The diversity of tools for studying fish movement across multiple years and various spatial scales gives researchers new abilities to address key science questions and management needs. Addressing these needs has the potential to improve upon existing fisheries management practices within the complexity of multi-jurisdictional governance in the Laurentian Great Lakes.     

Identifying sources and year classes contributing to invasive grass carp in the Laurentian Great Lakes

Relative contributions of aquaculture-origin and naturally-reproduced grass carp (Ctenopharyngodon idella) in the Laurentian Great Lakes have been unknown. We assessed occurrence and distribution of aquaculture-origin and wild grass carp in the Great Lakes using ploidy and otolith stable oxygen isotope (δ¹⁸O) data. We inferred natal river and dispersal from natal location for wild grass carp using otolith microchemistry and estimated ages of wild and aquaculture-origin fish to infer years in which natural reproduction and introductions occurred. Otolith δ¹⁸O indicated that the Great Lakes contain a mixture of wild grass carp and both diploid and triploid, aquaculture-origin grass carp. Eighty-eight percent of wild fish (n = 49 of 56) were caught in the Lake Erie basin. Otolith microchemistry indicated that most wild grass carp likely originated in the Sandusky or Maumee rivers where spawning has previously been confirmed, but results suggested recruitment from at least one other Great Lakes tributary may have occurred. Three fish showed evidence of movement between their inferred natal river in western Lake Erie and capture locations in other lakes in the Great Lakes basin. Age estimates indicated that multiple year classes of wild grass carp are present in the Lake Erie basin, recruitment to adulthood has occurred, and introductions of aquaculture-origin fish have happened over multiple years. Knowledge of sources contributing to grass carp in the Great Lakes basin will be useful for informing efforts to prevent further introductions and spread and to develop strategies to contain and control natural recruitment.     

Polymethylene-interrupted fatty acids: Biomarkers for native and exotic mussels in the Laurentian Great Lakes

Freshwater organisms synthesize a wide variety of fatty acids (FAs); however, the ability to synthesize and/or subsequently modify a particular FA is not universal, making it possible to use certain FAs as biomarkers. Herein we document the occurrence of unusual FAs (polymethylene-interrupted fatty acids; PMI-FAs) in select freshwater organisms in the Laurentian Great Lakes. We did not detect PMI-FAs in: (a) natural seston from Lake Erie and Hamilton Harbor (Lake Ontario), (b) various species of laboratory-cultured algae including a green alga (Scenedesmus obliquus), two cyanobacteria (Aphanizomenon flos-aquae and Synechococystis sp.), two diatoms (Asterionella formosa, Diatoma elongatum) and a chrysophyte (Dinobryon cylindricum) or, (c) zooplankton (Daphnia spp., calanoid or cyclopoid copepods) from Lake Ontario, suggesting that PMI-FAs are not substantively incorporated into consumers at the phytoplankton-zooplankton interface. However, these unusual FAs comprised 4-6% of total fatty acids (on a dry tissue weight basis) of native fat mucket (Lampsilis siliquoidea) and plain pocketbook (L. cardium) mussels and in invasive zebra (Dreissena polymorpha) and quagga (D. bugensis) mussels. We were able to clearly partition Great Lakes' mussels into three separate groups (zebra, quagga, and native mussels) based solely on their PMI-FA profiles. We also provide evidence for the trophic transfer of PMI-FAs from mussels to various fishes in Lakes Ontario and Michigan, further underlining the potential usefulness of PMI-FAs for tracking the dietary contribution of mollusks in food web and contaminant-fate studies.     

A systematic review of the literature on plastic pollution in the Laurentian Great Lakes and its effects on freshwater biota

Plastic pollution is ubiquitous in freshwater systems worldwide, and the Laurentian Great Lakes are no exception. We conducted a systematic review to synthesize the current state of the literature on plastic pollution, including macroplastics (>5 mm) and microplastics (<5 mm), in the Great Lakes. Thirty-four publications were used in our systematic review. We found ubiquitous contamination of microplastics in surface water, with maximum abundances exceeding those in ocean gyres. There are also high levels of plastic contamination reported across benthic sediments and shorelines of the Great Lakes. Citizen science data reveals macroplastic across Great Lakes shorelines, with more than three million pieces of plastic litter recorded over a span of three years. We completed a second systematic review of plastic pollution and its impact on freshwater ecosystems in general to inform how plastic in the Great Lakes may impact wildlife. Among studies published in the literature, we found 390 tested effects, 234 (60%) of which were detected and 156 (40%) of which were not; almost all of the freshwater effects (>98%) were tested on microplastics. Based on a subset of these papers, we found that the shape and size of a particle likely affects whether an effect is detected, e.g., more effects are detected for smaller particles. Finally, we identify gaps in scientific knowledge that need to be addressed and discuss how the state of the science can inform management strategies.     

The flathead catfish invasion of the Great Lakes

A detailed review of historical literature and museum data revealed that flathead catfish were not historically native in the Great Lakes Basin, with the possible exception of a relict population in Lake Erie. The species has invaded Lake Erie, Lake St. Clair, Lake Huron, nearly all drainages in Michigan, and the Fox/Wolf and Milwaukee drainages in Wisconsin. They have not been collected from Lake Superior yet, and the temperature suitability of that lake is questionable. Flathead catfish have been stocked sparingly in the Great Lakes and is not the mechanism responsible for their spread. A stocking in 1968 in Ohio may be one exception to this. Dispersal resulted from both natural range expansions and unauthorized introductions. The invasion is ongoing, with the species invading both from the east and the west to meet in northern Lake Michigan. Much of this invasion has likely taken place since the 1990s. This species has been documented to have significant impacts on native fishes in other areas where it has been introduced; therefore, educating the public not to release them into new waters is important. Frequent monitoring of rivers and lakes for the presence of this species would detect new populations early so that management actions could be utilized on new populations if desired.     

Heterogeneity of bacterial communities within the zebra mussel (Dreissena polymorpha) in the Laurentian Great Lakes Basin

We analyzed and compared the structure of bacterial communities associated with zebra mussel mantle cavity fluid, gills, and gut samples collected from Lake Loon, an inland lake in Michigan's Lower Peninsula (U.S.A.) using partial 16S rRNA gene sequencing. A total of 713 cloned 16S ribosomal gene sequences were checked for similarity to existing 16S sequences in two public databases: the Ribosomal Database Project and BLAST. Based on a 98% sequence similarity threshold, a total of 355 phylotypes belonging to 12 bacterial phyla and the phylum Bacillariophyta (diatoms) were identified in zebra mussel samples. A dominance of sequences belonging to the class γ-proteobacteria was observed in the mantle cavity clone libraries (P < 0.0001). Significant sample-specific sequence associations (P < 0.001) included members of the orders Pseudomonadales and Vibrionales in mantle cavity fluid and gut clone libraries, members of both the phylum Actinobacteria and the class δ-proteobacteria in gill clone libraries, and the Cyanobacteria/Bacillariophyta group in gut clone libraries. Furthermore, our results suggest that the zebra mussel may serve as a reservoir for facultative and opportunistic pathogenic bacteria, e.g., Clostridium spp., Flavobacterium spp. and Mycobacterium spp., for many aquatic and terrestrial animals. This work constitutes the first account of the heterogeneity of bacterial communities associated with multiple compartments within the zebra mussel. The information gained in this study significantly contributes to what is known regarding the microbial ecology of the zebra mussel and its role in disease ecology and food-web shifts in the Great Lakes ecosystem.     

Risk-based classification and interactive map of watersheds contributing anthropogenic stress to Laurentian Great Lakes coastal ecosystems

We describe development anthropogenic stress indices for coastal margins of the Laurentian Great Lakes basin. Indices were derived based on the response of species assemblages to watershed-scale stress from agriculture and urbanization. Metrics were calculated for five groups of wetland biota: diatoms, wetland vegetation, aquatic invertebrates, fishes, and birds. Previously published community change points of these assemblages were used to classify each watershed as ‘least-disturbed’, ‘at-risk’, or ‘degraded’ based on community response to these stressors. The end products of this work are an on-line map utility and downloadable data that characterize the degree of agricultural land use and development in all watersheds of the US and Canadian Great Lakes basin. Discrepancies between the observed biological condition and putative anthropogenic stress can be used to determine if a site is more degraded than predicted based on watershed characteristics, or if remediation efforts are having beneficial impacts on site condition. This study provides a landscape-scale evaluation of wetland condition that is a critical first step for multi-scale assessments to help prioritize conservation or restoration efforts.     

Toxaphene trends in the Great Lakes fish

As part of the U.S. Great Lakes Fish Monitoring and Surveillance Program (GLFMSP), more than 300 lake trout (Salvelinus namaycush) and walleye (Stizostedion vitreum vitreum) collected from the Laurentian Great Lakes each year from 2004 to 2009, have been analyzed for total toxaphene and eight selected congeners. The analytical results show fish toxaphene concentrations are quite different among lakes. Between 2004 and 2009, Lake Superior lake trout had the highest concentration (119 to 482 ng/g) and Lake Erie walleye had the lowest concentration (18 to 47 ng/g). Combining these results with the historical total toxaphene data (1977–2003), temporal changes were examined for each lake. Because of different analytical methods used in the previous studies, the historical data were adjusted using a factor of 0.56 based on a previous inter-method comparison in our laboratory. Trend analysis using an exponential decay regression showed that toxaphene in Great Lakes fish exhibited a significant decrease in all of the lakes with t_1/2 (confidence interval) of 0.9 (0.8–1.1) years for Lake Erie walleye, 3.8 (3.5–4.1) years for Lake Huron lake trout, 5.6 (5.1–6.1) years for Lake Michigan lake trout, 7.5 (6.7–8.4) years for Lake Ontario lake trout and 10.1 (8.2–13.2) years for Lake Superior lake trout. Parlars 26, 50 and 62 were the dominant toxaphene congeners accounting for 0.53% to 41.7% of the total toxaphene concentration. Concentrations of these congeners generally also decreased over time.     

The benthic community of the Laurentian Great Lakes: Analysis of spatial gradients and temporal trends from 1998 to 2014

We used the results of seventeen years of Great Lakes benthic monitoring conducted by the U.S. EPA's Great Lakes National Program Office to describe the spatial and temporal patterns of benthic communities, assess their status, trends, and main drivers, and to infer the potential impact of these community changes on ecosystem functioning. Benthic abundance and diversity were higher at shallow (<70?m in depth) stations with chlorophyll concentrations above 3?μg/L than at deeper sites (<1?μg/L). We infer that lake productivity, measured by chlorophyll was likely the major driver of benthic abundance and diversity across lakes. Consequently, benthic diversity and abundance were the highest in the most productive Lake Erie, followed by lakes Ontario, Michigan, Huron, and Superior. Multivariate analysis distinguished three major communities shared among lakes (littoral, sublittoral, and profundal) that differed in species composition and abundance, functional group diversity, and tolerance to organic pollution. Analysis of temporal trends revealed that the largest changes occurred in profundal communities, apparent in significant shifts in dominant taxa across all lakes except Lake Superior. In lakes Michigan, Huron, and Ontario, the former dominant Diporeia was replaced with Dreissena and Oligochaeta. Profundal species, with the exception of dreissenids, became less abundant, and their depth distribution has shifted. In contrast, density and diversity of native littoral and sublittoral communities increased. The invasion of dreissenids was among the most important drivers of changes in benthic communities. Continued monitoring is critical for tracking unprecedented changes occurring in the Great Lakes ecosystem.     

Great Lakes chloride trends: Long-term mass balance and loading analysis

Surveillance data collected over the past 150 years are compiled and analyzed to identify chloride trends in the Laurentian Great Lakes. These data indicate that chloride levels started rising in the mid-19th century and began accelerating in the early twentieth century. Lake Superior's and Lake Michigan's concentrations have continued to increase steadily and currently stand at their maximum recorded levels. In contrast, lakes Huron, Erie and Ontario reached peak levels between 1965 and 1975, but then began to decline. However, recent data indicate that the chloride concentrations in these lakes are now increasing again. Because loading data are not readily available, a mass-balance model is employed to estimate the chloride inputs required to account for the concentration trends. This inverse analysis yields computed load reductions that are consistent with reported industrial load reductions during the last three decades of the 20th century. Hence, it appears that the improvements were for the most part attributable to industrial controls. The model is also used to predict that if loads are held fixed at 2006 levels, concentrations in all lakes will continue to increase with the most dramatic rise occurring in Lake Michigan which will ultimately approach the level of Lake Erie.     

Historical changes and current status of crayfish diversity and distribution in the Laurentian Great Lakes

Despite increasing recognition of the importance of invertebrates, and specifically crayfish, to nearshore food webs in the Laurentian Great Lakes, past and present ecological studies in the Great Lakes have predominantly focused on fishes. Using data from many sources, we provide a summary of crayfish diversity and distribution throughout the Great Lakes from 1882 to 2008 for 1456 locations where crayfish have been surveyed. Sampling effort was greatest in Lake Michigan, followed by lakes Huron, Erie, Superior, and Ontario. A total of 13 crayfish species occur in the lakes, with Lake Erie having the greatest diversity (n = 11) and Lake Superior having the least (n = 5). Five crayfish species are non-native to one or more lakes. Because Orconectes rusticus was the most widely distributed non-native species and is associated with known negative impacts, we assessed its spread throughout the Great Lakes. Although O. rusticus has been found for over 100 years in Lake Erie, its spread there has been relatively slow compared to that in lakes Michigan and Huron, where it has spread most rapidly since the 1990s and 2000, respectively. O. rusticus has been found in both lakes Superior and Ontario for 22 and 37 years, respectively, and has expanded little in either lake. Our broad spatial and temporal assessment of crayfish diversity and distribution provides a baseline for future nearshore ecological studies, and for future management efforts to restore native crayfish and limit non-native introductions and their impact on food web interactions.     

Seasonal patterns in hydrochemical mixing in three Great Lakes rivermouth ecosystems

Rivermouth ecosystems in the Laurentian Great Lakes represent complex hydrologic mixing zones where lake and river water combine to form biologically productive areas that are functionally similar to marine estuaries. As urban, industrial, shipping, and recreational centers, rivermouths are the focus of human interactions with the Great Lakes and, likewise, may represent critical habitat for larval fish and other biota. The hydrology and related geomorphology in these deltaic systems form the basis for ecosystem processes and wetland habitat structure but are poorly understood. To this end, we examined hydrogeomorphic structure and lake-tributary mixing in three rivermouths of intermediate size using water chemistry, stable isotopes, and current profiling over a five-month period. In rivermouths of this size, the maximum depth of the rivermouth ecosystem influenced water mixing, with temperature-related, density-dependent wedging and layering that isolated lake water below river water occurring in deeper systems. The inherent size of the rivermouth ecosystem, local geomorphology, and human modifications such as shoreline armoring and dredging influenced mixing by altering the propensity for density differences to occur. The improved scientific understanding and framework for characterizing hydrogeomorphic processes in Great Lakes rivermouths across a disturbance gradient is useful for conservation, management, restoration, and protection of critical habitats needed by native species.     

Impacts on biodiversity and species changes in the lower Great Lakes

The Great Lakes basin ecosystem evolved after the retreat of the last ice sheet, about 10 000 years ago. Today, the complex of species present in the Great Lakes and much of the visible landscape bears little resemblance to that found some 400 years ago. Rather, the effects of various aspects of human development have caused major changes in the natural biodiversity. Lessons learned in the lower Great Lakes are applicable to many lakes around the world that have undergone agricultural, industrial and urban development in their drainage basins and have become managed, artificial ecosystems.     

Revisiting the Great Lakes Water Quality Agreement phosphorus targets and predicting the trophic status of Lake Michigan

LM3-Eutro is a high-resolution eutrophication model with several improved features lacking in historical Great Lakes models. We calibrated LM3-Eutro using a 2-year (1994-1995) dataset and performed a hindcast simulation from 1976 to 1995 to evaluate the model's ability to make predictions over an extended period of time. Results show a reasonable agreement between model output and field data over this time period. The model predicted that an annual loading of 5600 metric tons (MT) would result in a lake-wide annual total phosphorus (TP) concentration of 7.5 μg L^− 1. Using best estimates of future TP loadings, LM3-Eutro forecasts suggest that Lake Michigan will remain oligotrophic and will continue to meet the 7 μg L^− 1 spring TP concentration Great Lakes Water Quality Agreement objective.     

European frogbit (Hydrocharis morsus-ranae) invasion facilitated by non-native cattails (Typha) in the Laurentian Great Lakes

Plant-to-plant facilitation is important in structuring communities, particularly in ecosystems with high levels of natural disturbance, where a species may ameliorate an environmental stressor, allowing colonization by another species. Increasingly, facilitation is recognized as an important factor in invasion biology. In coastal wetlands, non-native emergent macrophytes reduce wind and wave action, potentially facilitating invasion by floating plants. We tested this hypothesis with the aquatic invasive species European frogbit (Hydrocharis morsus-ranae; EFB), a small floating plant, and invasive cattail (Typha spp.), a dominant emergent, by comparing logistic models of Great Lakes-wide plant community data to determine which plant and environmental variables exerted the greatest influence on EFB distribution at multiple scales. Second, we conducted a large-scale field experiment to evaluate the effects of invasive Typha removal treatments on an extant EFB population. Invasive Typha was a significant predictor variable in all AIC-selected models, with wetland zone as the other most common predictive factor of EFB occurrence. In the field experiment, we found a significant reduction of EFB in plots where invasive Typha was removed. Our results support the hypothesis that invasive Typha facilitates EFB persistence in Great Lakes coastal wetlands, likely by ameliorating wave action and wind energy. The potential future distribution of EFB in North America is vast due in part to the widespread and expanding distribution of invasive Typha and other invading macrophytes, and their capacity to facilitate EFB's expansion, posing significant risk to native species diversity in Great Lakes coastal wetlands.     

A pound of prevention,plus a pound of cure: Early detection and eradication of invasive species in the Laurentian Great Lakes

Ballast water regulations implemented in the early 1990s appear not to have slowed the rate of new aquatic invasive species (AIS) establishment in the Great Lakes. With more invasive species on the horizon, we examine the question of whether eradication of AIS is a viable management strategy for the Laurentian Great Lakes, and what a coordinated AIS early detection and eradication program would entail. In-lake monitoring would be conducted to assess the effectiveness of regulations aimed at stopping new AIS, and to maximize the likelihood of early detection of new invaders. Monitoring would be focused on detecting the most probable invaders, the most invasion-prone habitats, and the species most conducive to eradication. When a new non-native species is discovered, an eradication assessment would be conducted and used to guide the management response. In light of high uncertainty, management decisions must be robust to a range of impact and control scenarios. Though prevention should continue to be the cornerstone of management efforts, we believe that a coordinated early detection and eradication program is warranted if the Great Lakes management community and stakeholders are serious about reducing undesired impacts stemming from new AIS in the Great Lakes. Development of such a program is an opportunity for the Laurentian Great Lakes resource management community to demonstrate global leadership in invasive species management.