Influence of lake levels on water extent,interspersion, and marsh birds in Great Lakes coastal wetlands

Coastal wetlands in the Laurentian Great Lakes undergo frequent, sometimes dramatic, physical changes at varying spatial and temporal scales. Changes in lake levels and the juxtaposition of vegetation and open water greatly influence biota that use coastal wetlands. Several regional studies have shown that changes in vegetation and lake levels lead to predictable changes in the composition of coastal wetland bird communities. We report new findings of wetland bird community changes at a broader scale, covering the entire Great Lakes basin. Our results indicate that water extent and interspersion increased in coastal wetlands across the Great Lakes between low (2013) and high (2018) lake-level years, although variation in the magnitude of change occurred within and among lakes. Increases in water extent and interspersion resulted in a general increase in marsh-obligate and marsh-facultative bird species richness. Species like American bittern (Botaurus lentiginosus), common gallinule (Gallinula galeata), American coot (Fulica americana), sora (Porzana carolina), Virginia rail (Rallus limicola), and pied-billed grebe (Podilymbus podiceps) were significantly more abundant during high water years. Lakes Huron and Michigan showed the greatest increase in water extent and interspersion among the five Great Lakes while Lake Michigan showed the greatest increase in marsh-obligate bird species richness. These results reinforce the idea that effective management, restoration, and assessment of wetlands must account for fluctuations in lake levels. Although high lake levels generally provide the most favorable conditions for wetland bird species, variation in lake levels and bird species assemblages create ecosystems that are both spatially and temporally dynamic.     

Great Lakes coastal wetlands as suitable habitat for invasive mute swans in Michigan

Great Lakes coastal wetlands provide critical habitat and food resources for more species than any other Great Lakes ecosystem. Due to past and current anthropogenic disturbances, coastal wetland area has been reduced by >50% while remaining habitat is frequently degraded. Invasive mute swans have contributed to the degradation of coastal wetlands by removing submergent vegetation and competitively excluding native species from breeding areas and food resources. Despite current control practices, mute swan population estimates in Michigan are ~8000, comparable to population estimates in the entire Atlantic Flyway of North America. We collected local abiotic data and adjacent land cover data at 3 scales from 51 sites during 2010 and 2011 and conducted 2 mute swan detection surveys each year during the summer and fall. We developed a single-species, single-season occupancy-based habitat suitability model to determine current and potential mute swan habitat among Great Lakes coastal wetlands. We found mute swans occupied heterotrophic coastal wetlands adjacent to urban areas, which were high in ammonium and oxidation-reduction potential and low in nitrates, dissolved oxygen, and turbidity. Our model provides managers with a valuable tool for rapidly identifying mute swan habitat areas for control efforts, particularly the need for targeting mute swan populations in or near urbanized areas. Our model will also aid managers in monitoring areas that mute swans may invade and prioritizing coastal wetland areas for restoration efforts.     

Water depth and lake-wide water level fluctuation influence on α- and β-diversity of coastal wetland fish communities

Coastal wetlands in the Laurentian Great Lakes are critical habitats for supporting fish diversity and abundance within the basin. Insight into the coupling of biodiversity patterns with habitat conditions may elucidate mechanisms shaping diverse communities. Within coastal wetlands, water depth as well as fluctuations in lake-wide water levels over inter-annual timescales, both have the potential to influence fish communities. Water level fluctuation can influence fish habitat structure (e.g., vegetation) in Great Lakes coastal wetlands, but it is unclear how water depth and lake-wide water level fluctuations affect fish community composition and diversity. Using β dissimilarity indices and multivariate ordination techniques, we assessed fish community structure among bulrush (Schoenoplectus acutus)-dominated wetlands in Saginaw Bay, Lake Huron, USA. We examined whether community structure was related to wetland water depth at the time of sampling and whether fish communities were more similar among years with similar Lake Huron water levels. Results suggested relatively high levels of both spatial (among wetlands) and temporal (among year) community dissimilarity that was driven primarily by species turnover. Thus, variability in water depths among wetlands and in Lake Huron water levels among years likely both contribute to regional fish diversity. Further, fish abundance and alpha diversity were positively correlated with wetland water depth at the time of sampling. Both climate change and anthropogenic water level stabilization may alter the magnitude and timing of water level fluctuations in the Great Lakes. Our data suggest that these changes could affect local fish community composition and regional fish diversity.     

Environmental predictors of phytoplankton chlorophyll-a in Great Lakes coastal wetlands

Coastal wetlands of the Laurentian Great Lakes are diverse and productive ecosystems that provide many ecosystem services, but are threatened by anthropogenic factors, including nutrient input, land-use change, invasive species, and climate change. In this study, we examined one component of wetland ecosystem structure – phytoplankton biomass – using the proxy metric of water column chlorophyll-a measured in 514 coastal wetlands across all five Great Lakes as part of the Great Lakes Coastal Wetland Monitoring Program. Mean chlorophyll-a concentrations increased from north-to-south from Lake Superior to Lake Erie, but concentrations varied among sites within lakes. To predict chlorophyll-a concentrations, we developed two random forest models for each lake – one using variables that may directly relate to phytoplankton biomass (“proximate” variables; e.g., dissolved nutrients, temperature, pH) and another using variables with potentially indirect effects on phytoplankton growth (“distal” variables; e.g., land use, fetch). Proximate and distal variable models explained 16–43% and 19–48% of variation in chlorophyll-a, respectively, with models developed for lakes Erie and Michigan having the highest amount of explanatory power and models developed for lakes Ontario, Superior, and Huron having the lowest. Land-use variables were important distal predictors of chlorophyll-a concentrations across all lakes. We found multiple proximate predictors of chlorophyll-a, but there was little consistency among lakes, suggesting that, while chlorophyll-a may be broadly influenced by distal factors such as land use, individual lakes and wetlands have unique characteristics that affect chlorophyll-a concentrations. Our results highlight the importance of responsible land-use planning and watershed-level management for protecting 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.     

A reference inventory for aquatic fauna of the Laurentian Great Lakes

The Laurentian Great Lakes encompass an expansive and diverse set of freshwater ecosystems that contain a concordantly large and diverse vertebrate and invertebrate fauna. Although numerous publications exist concerning the composition and distribution of this fauna, there is at present no single readily available resource that brings all this information together. Here, we present and describe the compilation process for a comprehensive Great Lakes aquatic fauna inventory covering fishes, reptiles, amphibians, zooplankton, mollusks, annelids, insects, mites, and various other aquatic invertebrates. Inventory entries were developed via an extensive search of literature and internet sources and are attributed with detailed nomenclature information, general lake and habitat occurrences, and supporting citations and links to life history and genetic marker information. The inventory scope is the Laurentian Great Lakes proper and their connecting rivers, and their fringing coastal wetlands and lower tributaries. Over 2200 unique taxa are contained in the inventory – 85% resolved to species and 14% to genus. The listing substantially expands previous richness estimates for invertebrates in the Great Lakes, but taxonomic resolution and spatial distribution information for them remains quite uneven. Example pattern analyses for fauna in this inventory show that aquatic vertebrates are generally more widely distributed than invertebrates, and that biodiversity is concentrated in the coastal margins. The inventory is being packaged into a public, searchable database that showcases the biodiversity of the Great Lakes aquatic fauna and can assist the research and management community in their biological investigations.     

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.     

Habitat Associations of Larval Fish in a Lake Superior Coastal Wetland

Information on the habitat associations of larval fishes in Great Lakes coastal wetlands (GLCW) is necessary to assist fisheries managers in the protection and management of critical habitats. Coastal wetlands serve as spawning grounds, nurseries, and forage areas for many important Great Lakes fish species. To determine the distribution of larval fish in coastal wetlands with regard to location and vegetation characteristics, we used a larval tow-sled to sample four macrohabitat types (sand-spit, inner and outer marsh, and river) across sparse, moderate, and dense vegetation densities (microhabitat) in Allouez Bay wetland near Lake Superior's western end. We captured 4,806 larval fish representing 16 species between May and August 1996. Allouez Bay is typical of other GLCW in species number and composition. The three most abundant species were spottail shiner (59% of the total catch), yellow perch (20% of total catch), and white sucker (10% of total catch). Significantly more fish and fish species (repeated-measures ANOVA) (p < 0.05) were caught at the sand-spit relative to the outer or inner marsh macrohabitats. Nearly all of the cyprinids and centrarchids were caught at the sand-spit habitat primarily in dense vegetation, while the majority of white suckers and trout-perch were caught in the river in sparse or moderate vegetation. Our study provides evidence for species-specific macrohabitat and microhabitat associations of larval fish in coastal wetlands. We suggest these associations are largely determined by adult spawning requirements and life-history strategies.     

Great Lakes coastal fish habitat classification and assessment

Basin-scale assessment of fish habitat in Great Lakes coastal ecosystems would increase our ability to prioritize fish habitat management and restoration actions. As a first step in this direction, we identified key habitat factors associated with highest probability of occurrence for several societally and ecologically important coastal fish species as well as community metrics, using data from the Great Lakes Aquatic Habitat Framework (GLAHF), Great Lakes Environmental Indicators (GLEI) and Coastal Wetland Monitoring Program (CWMP). Secondly, we assessed whether species-specific habitat was threatened by watershed-level anthropogenic stressors. In the southern Great Lakes, key habitat factors for determining presence/absence of several species of coastal fish were chlorophyll concentrations, turbidity, and wave height, whereas in the northern ecoprovince temperature was the major habitat driver for most of the species modeled. Habitat factors best explaining fish richness and diversity were bottom slope and chlorophyll a. These models could likely be further improved with addition of high-resolution submerged macrophyte complexity data which are currently unavailable at the basin-wide scale. Proportion of invasive species was correlated primarily with increasing maximum observed inorganic turbidity and chlorophyll a. We also demonstrate that preferred habitat for several coastal species and high-diversity areas overlap with areas of high watershed stress. Great Lakes coastal wetland fish are a large contributor to ecosystem services as well as commercial and recreational fishery harvest, and scalable basin-wide habitat models developed in this study may be useful for informing management actions targeting specific species or overall coastal fish biodiversity.     

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.     

Relationship of road density and marsh condition to turtle assemblage characteristics in the Laurentian Great Lakes

Human-induced degradation of coastal wetlands often leads to altered trophic dynamics and species assemblages. Here we use data from 77 coastal marshes in three Laurentian Great Lakes collected between 2001 and 2007 to examine the relationship between human disturbance (road density and wetland quality) and characteristics of aquatic turtle assemblages, including species richness and abundances. Painted turtles (Chrysemys picta) were encountered disproportionately in degraded wetlands and the probability of occurrence decreased with improved site quality. Abundance of painted turtles peaked, however, at intermediate road density in surrounding 1- and 2-km buffers. Across all sites, species richness was highest and common snapping turtles (Chelydra serpentina) were most abundant in wetlands with intermediate water quality. The common musk turtle (Sternotherus odoratus) was absent from degraded wetlands in the lower lakes (Erie and Ontario) that fell within their historical range, but reached high abundances in marshes of Georgian Bay and the North Channel, a region with relatively low human disturbance. Analysis of sex ratios in painted turtles revealed a significant male bias in an area with high road density, while the sex ratio did not differ significantly from 1:1 in a less developed region, consistent with reports of high female mortality in urban areas.     

Exotic and Invasive Aquatic Plants in Great Lakes Coastal Wetlands: Distribution and Relation to Watershed Land Use and Plant Richness and Cover

We use data from inundated-area surveys of 58 coastal wetlands spanning a gradient of anthropogenic impacts across all five Laurentian Great Lakes to describe the distribution of nine exotic and invasive taxa of aquatic plants. We found plants that were exotic or have invasive strains to be substantially more prevalent in wetlands in Lakes Erie and Ontario than in Lakes Superior and Huron, with Lake Michigan wetlands intermediate. Najas minor (slender naiad), Butomus umbellatus (flowering rush), and Hydrocharis morsus-ranae (European frogbit) were restricted to the lower lakes and rarely dominant. Myriophyllum spicatum (Eurasian milfoil), Potamogeton crispus (curly pondweed), Lythrum salicaria (purple loosestrife), Phalaris arundinacea (reed canary grass), Phragmites australis (common reed), and Typha sp. (cattail) were more widespread and except for P. crispus, often among the dominant taxa. None of the submerged or floating-leaf exotic taxa were associated with altered total plant cover or richness, although M. spicatum, P. crispus, and native Stuckenia pectinatus (sago pondweed) were positively associated with agricultural intensity in the watershed (a surrogate for nutrient loading). Emergent P. australis, L. salicaria, and Typha were more likely to be present and dominant as agricultural intensity increased, and were associated with elevated emergent cover and decreased emergent genera richness. Effects of dominant taxa on plant cover and richness were readily detected using ordinal data from 100 m inundated segments but were harder to discern with data aggregated to the wetland scale. The sum of shoreline-wide abundance scores for four easily identified taxa (S. pectinata, P. australis, Typha, and L. salicaria) is proposed as a rapidly-measured indicator of anthropogenic disturbance across the Great Lakes.     

Invasive Plant Species in Diked vs. Undiked Great Lakes Wetlands

We compared the standing vegetation, seed banks, and substrate conditions in seven pairs of diked and undiked wetlands near the shores of Lake Michigan and Lake Huron, North America. Our analysis tested the null hypothesis that construction of artificial dikes has no effect on the vulnerability of Great Lakes coastal wetlands to non-native and native invasive species. Both the standing vegetation and seed banks in diked wetlands contained significantly more species and individuals of invasive plants. In addition, diked wetlands exhibited significantly higher levels of organic matter and nutrient levels, and significantly higher average pH. Two pervasive non-native invasive species in the Great Lakes region, Lythrum salicaria (purple loosestrife) and Phalaris arundinacea (reed canary grass) were significantly more abundant in diked wetlands. Typha spp. (cattail) also formed a much higher percent vegetation cover in the diked wetlands. Our results support the view that diking of shoreline wetlands modifies natural hydrologic regimes, leading to nutrient-rich aquatic environments that are vulnerable to invasion. The shallower, more variable water levels in non-diked wetlands, on the other hand, appear to favor another undesirable invasive species, Phragmites australis (common reed grass).     

Relative importance of macrophyte community versus water quality variables for predicting fish assemblages in coastal wetlands of the Laurentian Great Lakes

Fish have been shown to be sensitive indicators of environmental quality in Great Lakes coastal wetlands. Fish composition also reflects aquatic macrophyte communities, which provide them with critical habitat. Although investigators have shown that the relationship between water quality and fish community structure can be used to indicate wetland health, we speculate that this relationship is a result of the stronger, more direct relationship between water quality and macrophytes, together with the ensuing interconnection between macrophyte and fish assemblages. In this study, we use data collected from 115 Great Lakes coastal marshes to test the hypothesis that plants are better predictors of fish species composition than is water quality. First we use canonical correspondence analysis (CCA) to conduct an ordination of the fish community constrained by water quality parameters. We then use co-correspondence analysis (COCA) to conduct a direct ordination of the fish community with the plant community data. By comparing the statistic ‘percent fit,’ which refers to the cumulative percentage variance of the species data, we show that plants are consistently better predictors of the fish community than are water quality variables in three separate trials: all wetlands in the Great Lakes basin (whole: 21.2% vs 14.0%; n = 60), all wetlands in Lakes Huron and Superior (Upper: 20.3% vs 18.8%; n =  32), and all wetlands in Georgian Bay and the North Channel (Georgian Bay: 18% vs 17%; n =  70). This is the largest study to directly examine plant–fish interactions in wetlands of the Great Lakes basin.     

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.     

Invasive cattail reduces fish diversity and abundance in the emergent marsh of a Great Lakes coastal wetland

Great Lakes coastal wetlands (GLCWs) provide critical fish habitat. The invasion of GLCWs by hybrid and narrow-leaved cattail, Typha × glauca and Typha angustifolia (hereafter Typha), homogenizes wetlands by out-competing native plant species and producing copious litter. However, the effect of this invasion on fish communities is little known. To measure the effect of Typha on fishes, we established plots in Typha invaded and native wetland emergent zones in a northern Lake Michigan coastal wetland, and measured environmental variables, plants, and fishes in each zone over two summers. Dissolved oxygen and water temperature were significantly lower in invaded compared to native plots. Invaded plots were dominated by Typha and its litter; whereas. sedges (Carex spp.) were the most abundant species in native plots. Fish abundance and species richness were significantly lower in Typha compared to native wetland plots. The Typha fish community was dominated by hypoxia tolerant mudminnow whereas other small, schooling, fusiform species such as cyprinids and fundulids were absent. These results illustrate the negative impact of a dominant invasive plant on Great Lakes fishes that is expected to be found in Typha invasions in other GLCWs.     

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.     

Reviewing uncertainty in bioenergetics and food web models to project invasion impacts: Four major Chinese carps in the Great Lakes

Bioenergetics and food web models are tools available for understanding and projecting the impacts of aquatic species invasions on food web structure and energy allocation of an ecosystem. However, uncertainty is inherent in modeling the impact of invasive species in novel ecosystems as assumptions must be made about physiological responses to novel environments and interactions with existing (native and non-native) species. Here we use the four major Chinese carps (FMCC) in the Laurentian Great Lakes as a case study to categorize and describe the suite of uncertainties inherent in projecting the impact of invasive species with bioenergetics and food web models. We approach this case study in a decision analytic framework, describing structural uncertainties, environmental variation, partial observability, partial controllability, and linguistic uncertainty. Finally, we review and give suggestions for how the use of methods including adaptive management, scenario planning, sensitivity analyses, and value of information as well as efforts to ensure clarity in language and model structure can enable modelers and managers to reduce and account for key uncertainties and make better decisions for the control of invasive species.