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16 November 2025

Ecological Restoration in Laurentian Great Lakes Wetlands: A Literature Review

,
and
1
Department of Physical and Environmental Sciences, Department of Biological Sciences, University of Toronto Scarborough, 1265 Military Trail, Toronto, ON M1C 1A4, Canada
2
Aquatic Research and Monitoring Section, Ministry of Natural Resources, DNA Building, Trent University, 2140 East Bank Drive, Peterborough, ON K9L 1Z8, Canada
*
Author to whom correspondence should be addressed.
Diversity2025, 17(11), 797;https://doi.org/10.3390/d17110797 
(registering DOI)
This article belongs to the Special Issue Ecological Restoration, Functioning and Conservation of Coastal Wetlands
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Abstract

The Laurentian Great Lakes basin, the largest surface freshwater ecosystem in the world, is heavily impacted by anthropogenic stressors including wetland degradation and loss. Ecological restoration has been identified as a tool to conserve and manage Great Lakes wetlands where studies undergo planning, action, and evaluation stages. A review of 57 studies (1967–2023) on Great Lakes coastal and inland wetlands was conducted to determine when and where ecological restoration is occurring, what approaches are being taken, and how effective they are. Restoration occurred most in coastal wetlands located in the United States of America and Lake Erie. The most implemented monitoring designs were control-impact and before-after-control-impact designs. A common ecological objective of restoration was altering species composition for invasive species management. Studies targeting changes in biological communities integrated ecological theory well within the three stages of restoration. Variable restoration effectiveness was detected, where a mix of ecological objectives were targeted and monitored through many study designs. Future restoration efforts would benefit from greater financial and monitoring investments (especially during the planning stage), continued integration of ecological theory, development of lake-specific benchmarks to assess restoration success, and a collaborative approach that includes practitioner knowledge and Traditional Ecological Knowledge.

1. Introduction

The Laurentian Great Lakes basin is the largest surface freshwater ecosystem in the world, covering 2,441,106 km2 and containing 21% of the global surface fresh water. The ecosystem comprises five lakes, four located in both Canada and the United States of America (USA) (Lake Superior, Lake Huron, Lake Erie, Lake Ontario), and Lake Michigan entirely located in the USA. A diverse range of providing, provisioning, regulating, cultural, and supporting ecosystem services are experienced by the ~35 million people living in the Great Lakes basin. As such, the ecosystem has great intrinsic, economic, cultural, social, and ecological value (e.g., [,]).
The Great Lakes basin has experienced extensive wetland loss since European settlement. On the Canadian side of the Great Lakes basin, it is estimated that 72% of wetlands present before settlement have been lost as of 2002 in southern Ontario [], with more recent estimates showing a loss of over 1.5 million hectares between 1800 and 2019 []. Historical aerial imagery of Lake Erie coastal wetlands from 1935 to 1980 showed that 80% total area of wetlands that existed prior to 1940 were lost by 1980 [] and comparisons between 1940 and 2003 showed that agricultural area continues to dominate land cover with more recent land conversion due to commercial, industrial, and residential development []. A similar story unfolds on the American side of the Great Lakes basin, where some states lost upwards of 85% of wetland habitat post settlement [], with more recent literature reporting wetland habitat losses of 50% across the basin [].
Wetland loss is projected to continue under the stressors of urbanization, climate change, and invasive species. Wetland habitat continues to be indirectly replaced with alternatives, such as stormwater-management ponds, that are often smaller in size and do not provide the same ecological functions and services of a natural wetland []. Climate-change impacts are also likely to synergistically interact with invasive species by lowering the resilience of wetlands and increasing the likelihood of invasion by non-native aquatic species []. In addition to the loss of habitat, existing wetland habitat in the Great Lakes is often fragmented, has lower native species richness [], and is under increasing threat from largescale anthropogenic stressors. The World Wildlife Fund Living Planet Report [] indicated that freshwater ecosystems are disproportionately impacted by anthropogenic stress, resulting in an 83% decline in population abundances of freshwater species between 1970 and 2018. Dudgeon et al. [] identified the five major threats impacting freshwater biodiversity as overexploitation, destruction or degradation of critical habitat, pollution, modification of flow regimes, and non-native species invasion. Reid et al. [] expanded this list to include 12 major stressors including climate change.
Despite the large-scale loss and destruction of wetland habitat, existing coastal and inland wetlands in the Great Lakes basin are valuable for their provision of ecosystem functions and services. Great Lakes wetlands have long-been recognized as the kidneys of the Great Lakes ecosystem because they serve as metabolic regulators that filter, recycle, and release nutrients and organic materials [,]. As highly dynamic heterogenous zones connecting terrestrial and aquatic ecosystems, wetlands serve as habitat for a wide range of terrestrial and aquatic organisms, often being identified as unique biodiversity hotspots across spatial and temporal scales (e.g., [,,]). Furthermore, coastal wetlands are natural barriers that protect coastal areas from flooding and erosion []. The conservation and restoration of wetlands have been identified as optimal nature-based solutions for counteracting the loss of biodiversity and improving the future resilience of freshwater ecosystems []. Lastly, wetlands are important landscape components to combat the effects of climate change, where the restoration of wetlands has been identified as the most important restoration activity occurring in North America [].
Ecological restoration refers to actions taken to assist degraded, damaged, or destroyed ecosystems with their recovery []. Implementing and evaluating the effectiveness of ecological restoration is often separated into a planning stage, an action stage, and an evaluation stage. In the Great Lakes basin, stages of ecological restoration are outlined in the frameworks of binational efforts towards ecosystem recovery, such as Remedial Action Plans (RAP) []. The primary stage of a RAP assesses an ecosystem within a defined area to identify beneficial-use impairments or changes within an ecosystem that negatively impact ecosystem functions or services. The second stage of the RAP identifies remediation actions that can address impairments to ecosystem functioning. The third stage requires evaluation to provide evidence that ecological restoration has resulted in recovery. RAP stages also integrate the International Union for Conservation of Nature (IUCN) standards guiding international restoration efforts with the first stage including practices of assessment, the second stage considering planning and design, and the final stage integrating monitoring and evaluation components [].
Given the value of, and current threats to, freshwater ecosystems, and the need to restore and protect the chemical, physical, and biological integrity of the Great Lakes basin under the Great Lakes Water Quality Agreement [], there is a well-established need to invest in monitoring, protecting, and recovering Great Lakes wetlands. Currently, efforts to synthesize approaches taken to restore Great Lakes wetlands and evaluate whether restoration actions have achieved their ecological outcomes are limited. This limitation is reflected in the primary literature where calls to action for evidence-based restoration [] and adaptive management plans [] have been made. Specifically, the need to evaluate the effectiveness of common habitat restoration practices relative to their intended ecological outcomes has been identified []. The overall objective of this study was to characterize the undertaking and evaluation of ecological restoration actions, including all human-mediated actions taken to protect, conserve, manage, or recover Great Lakes wetland ecosystems. A literature review was done to answer the following questions:
  • When and where has ecological restoration occurred in Great Lakes wetlands?
  • What approaches were taken to implement and evaluate ecological restoration in Great Lakes wetlands?
  • How effective was ecological restoration in Great Lakes wetlands?

2. Materials and Methods

To review the published literature on ecological restoration of Great Lakes coastal and inland wetlands, the following terms were searched in the Web of Science core collection on 8 November 2023, producing 2166 results:
  • Great Lakes OR Lake Superior OR Lake Erie OR Lake Ontario OR Lake Huron OR Lake Michigan
And
  • Wetland OR swamp OR marsh OR fen OR bog
And
  • Restore OR rehabilitate OR remediate OR mitigate OR conserve OR improve OR augment OR revitalize OR naturalize OR enhance OR construct OR create OR revamp OR repair OR fix OR refurbish OR remodel OR recover OR reclaim OR renew OR revive OR rejuvenate OR alter OR manage
A secondary search with the same terms was conducted on 16 April 2024 using Google Scholar to evaluate the efficacy of the primary search []. The first 200 papers, sorted by relevance, were reviewed. Because only two relevant papers were identified in the first 200 papers and several relevant papers discovered in the primary search were also present in the Google Scholar search, a stopping point was deemed justified as per guidelines provided by the Collaboration for Environmental Evidence []. All papers from both searches were reviewed based on the abstract to determine whether papers would be retained for full-paper analysis (Figure 1, Appendix A Table A1). Ten papers were initially reviewed by two authors (DR and SMR) to evaluate coding criteria, and the remaining papers were reviewed by DR. Given the shift in the restoration field from targeting individual species recovery to targeting ecosystem recovery and pressing international targets for the conservation of biodiversity (i.e., Global Biodiversity Framework and the United Nation’s Decade on Ecosystem Restoration), the focus was ecosystem-based, ecological restoration, as opposed to species-specific restoration.
Figure 1. Flow diagram outlining the searching and screening process for a literature review on ecological restoration actions in Laurentian Great Lakes coastal and inland wetlands.
To determine when and where ecological restoration occurred, publication year, country, and lake where the actions occurred were classified for each paper. Wetland type was defined for each study as either a coastal wetland or an inland wetland. Freshwater coastal wetlands are influenced by Great Lakes hydrology and located along the Great Lakes shoreline or an adjacent lake (e.g., Muskegon Lake) or river (e.g., St. Lawrence and Detroit rivers). Inland wetlands are physically separated from the Great Lakes and are located inland. The ecological restoration of inland wetlands, located in upper watersheds of the basin, is an important activity inherently tied to improving the health of the Great Lakes ecosystem as a whole. The scale (in hectares) and maximum number of years studied after implementation were recorded for each study. To summarize ecological restoration practices, the terms used to describe restoration actions were recorded. Papers were grouped by common ecological restoration actions taken and common ecological objectives targeted. Actions were classified using the following categories: managing invasive species; reversing agricultural land; managing hydrology; and, creating wetland habitat. For some studies, multiple actions were identified. Ecological objectives were classified using the following categories: altering species composition; altering species composition and physical habitat; altering species composition and ecological function; altering sediment, nutrient, or contaminant loading and/or cycling; and, creating/modifying physical habitat.
The integration of ecological theory is an important component to planning, implementing, and evaluating restoration actions []. The integration of ecological theory can produce more complex study designs, often requiring greater financial investment and expertise. However, well-designed studies integrating ecological theory can diagnose the causes of restoration failure or success, where the accumulation of knowledge over time can be used to improve future restoration efforts. To evaluate how well restoration actions and evaluations incorporated ecological theory, 36 papers with a component of biological community analysis were further studied (e.g., []). Each paper was scored as one (present) or zero (not present) for whether ecological theory was used in the three stages of ecological restoration (planning, action, and evaluation). Studies based in ecological theory may test theories of succession/community assemblage, competition, alternative stable states, functional traits, and resilience, or test the effectiveness of specific mechanisms, such as improving connectivity or interspersion, in relation to theories (see Table 1 in []). For the first planning stage, the introduction section of the paper was examined to determine if ecological theory was present. For the action stage, if the objectives and methods sections of the paper tested ecological theory, the paper was scored as having ecological theory present. Lastly, the discussion section of the paper was analyzed for the presence of ecological theory in the third evaluation stage (Appendix A Table A2). Each paper received a cumulative score out of three, where papers with a score of three integrated ecological theory in all stages. Linkages among actions taken, ecological objectives targeted, and scores for the integration of ecological theory were visualized using an alluvial plot for all studies with a component of community analysis. All alluvial plots were created in R using the ggalluvial package [].
A well-designed monitoring program is an important component in the evaluation stage of restoration, where assessing the outcomes of ecological restoration is required []. Monitoring design for each study was defined as: descriptive (D)—actions analyzed without spatial or temporal controls; before-after (BA)—actions analyzed by comparing metrics before and after action at the site; control-impact (CI)—actions analyzed by comparing metrics at control and impact (action) sites; or before-after-control-impact (BACI)—actions analyzed by comparing metrics at control and impact (action) sites before and after action. The type of reference condition used for comparison was classified as historical, pristine conditions (>20 years prior to action implementation), current positive and naturalized conditions (<20 years prior to action implementation), current negative and disturbed conditions (<20 years prior to action implementation), or no condition for studies where no formal comparison to a reference condition was made [,,] (Appendix A Table A3). Statistical methods used were defined as univariate (e.g., t-tests, analyses of variance (ANOVA)), multivariate (e.g., ordination, correspondence, redundancy analyses), or summary statistics (no formal statistical analysis). Linkages among study design, reference condition, and statistical method were visualized using an alluvial plot. For each study, the type of variable monitored to evaluate the actions was recorded. Studies that monitored only physical habitat variables and/or ecosystem functions were defined as monitoring an abiotic response. Papers monitoring species composition and/or structural diversity had the response variable defined by the taxa that was evaluated. Papers evaluating a combination of abiotic and biotic responses, or multiple taxa, were classified as having multiple response variables. The effectiveness of each study was reported in relation to their ecological objective(s) as having: a positive effect, where the objective(s) was achieved or an effect was observed; a negative effect, where the objective(s) was not achieved or no effect were observed; or, a variable effect, where some, but not all objectives were achieved.
The definition of criteria for some cases was not straightforward, and some adaptations were made. Above- and below-ground biomass was considered an abiotic response because it was used to measure primary productivity. Several studies had both community- and species-specific analyses (e.g., monitoring young-of-the-year Northern Pike (Esox lucius) and community vegetation response), included both field and lab studies, or considered other land types that were not wetlands. In these cases, only the community-level data, field-based data, or wetland-specific results were included in the analysis, and the papers were classified based only on the relevant data.

3. Results

3.1. When and Where Has Ecological Restoration Occurred?

Fifty-seven papers were assessed as relevant for the study, mostly (except for two papers) discovered in the Web of Science search. Very few grey literature publications (i.e., government and technical reports, student theses, conference proceedings) were encountered during both searches: none of which met the inclusion criteria. Papers were published between 1967 and 2023, with higher numbers of papers published in the last 9 years (Figure 2). Eighty-one percent of studies (46 papers) were located in the USA, 18% (10) in Canada, and one paper was binational. Very few studies occurred in multiple lakes (5), and most studies occurred in lakes Erie (20), Michigan (19), and Ontario (14), and the fewest in lakes Huron (8) and Superior (5). Restoration actions were most often implemented in coastal (37) or inland (18) wetlands, with few studies occurring in both wetland ecosystem types (2). Actions were most often implemented in one wetland (33) and targeted ≤ 50 ha of wetland area (24). Sites were commonly evaluated one to two years after implementation, although a wide range of time (0–69 y) since implementation was studied (see Table S1 for a complete list of papers and their classifications).
Figure 2. Number of papers, by study type, published in each 5-year period that evaluated ecological restoration actions for coastal and inland wetlands in the Great Lakes basin (n = 57 published papers). Studies were classified as descriptive (D), before-after (BA), control-impact (CI), or before-after-control-impact (BACI) studies. One paper published in 1967 that implemented a BACI study design is not included in the figure.

3.2. What Approaches Were Taken to Implement and Evaluate Ecological Restoration?

The most common terms used to describe actions were restoration (30 papers) or management (13). Very few papers used terms such as mitigation, control, conservation practices, treatment, rehabilitation, cropping, enhancement, habitat modification, reconnection, or a combination thereof. The most common action was controlling invasive species, such as European Common Reed (Phragmites australis, herein referred to as Phragmites), hybrid Typha, or Common Carp (Cyprinus carpio). Additional actions included the conversion of agricultural land back to wetlands, creation of new wetlands, and alterations to hydrological processes, commonly including the reconnection of previously isolated wetland habitat. Many papers evaluated a combination of these actions. One paper evaluated wetland restoration, but the type of action could not be determined []. Actions were often done with the ecological objective of improving or altering the composition of biotic communities (26 papers). Fewer projects were done to manage nutrient, sediment, or contaminant loading and/or cycling (19), create habitat or improve existing wildlife habitat (1), alter or manage hydrological processes (1), or a combination thereof (10).
Thirty-six papers used community analysis in their study design. Most papers with community analysis integrated ecological theory within all three stages of restoration (22 papers), with fewer papers integrating ecological theory into only two stages (9), commonly the planning and assessment stages, or in one stage, commonly the assessment stage (5). When considering the links between actions, ecological objectives, and ecological theory, studies managing invasive species or creating wetland habitat with the goal of altering species composition were common and often integrated ecological theory in all stages of ecological restoration. Alternatively, studies evaluating agricultural reversal actions or multiple actions often addressed a wider range of ecological objectives. Lastly, hydrological management studies commonly attempted to alter species composition or nutrient and/or sediment loading and cycling (Figure 3).
Figure 3. Alluvial plot linking the type of restoration action taken, the intended ecological objectives of the action, the integration of ecological theory, and the effectiveness of the action in relation to the ecological objective, for papers evaluating ecological restoration in Great Lakes coastal and inland wetlands with a component of community analysis (n = 36). The ecological theory score was calculated by the cumulative number of phases (planning, action, and evaluation) ecological theory was integrated for each study. The vertical width of the coloured lines is proportional to the flow quantity (i.e., study counts).
Most studies compared control and impact sites (22 papers), with BACI studies being the second most common (18). Remaining studies were either descriptive (without any comparison to reference conditions) (10) or compared the site before and after the actions were implemented (7). Most papers used a current positive reference condition for comparison (24), with fewer papers using a negative reference condition (11) or both in combination (10). Only one paper each used historical positive conditions or all reference conditions for comparison (historical/current and positive/negative). Studies with a D or BA design commonly implemented univariate statistics, with the latter study design often comparing restoration conditions to a current negative reference condition. Studies with a BACI or CI design tended to utilize a wider range of statistical methodologies and reference conditions for comparison, where BACI studies commonly compared restoration conditions to current negative conditions. Papers with a CI study design tended to compare restoration conditions to a wide range of reference conditions and were the only study-type to use a historical reference condition (Figure 4).
Figure 4. Alluvial plot linking the study design, type of reference condition used for comparison, and statistical method implemented for papers evaluating ecological restoration in Great Lakes coastal and inland wetlands (n = 57). The vertical width of the coloured lines is proportional to the flow quantity (i.e., study counts).

3.3. How Effective Was Ecological Restoration?

Most studies monitored several measures of response (21 papers) or an abiotic response (19), such as water chemistry or quality, geochemical processes, nutrient processing, or carbon loading/cycling. Twelve papers monitored vegetation response, whereas only two papers monitored fish community response. Only one paper each monitored macroinvertebrates, turtles, or bacteria community response. Most papers reported varying effectiveness, with only very few identifying successful outcomes (Figure 5). Thirty-five papers had variable effects (i.e., different effects across monitoring indicators), often showing positive effects compared to negative controls, but negative effects compared to positive or natural controls. Variable effects were common when studies monitored multiple variables, indices, or taxa. When considering effectiveness in relation to the restoration actions and their ecological objectives, actions and objectives that targeted abiotic components of the ecosystem, such as creating or modifying habitat, hydrological management, or loading and cycling, were the only objectives that produced negative outcomes. Papers with limited integration of ecological theory (score of one), almost exclusively reported positive outcomes. Papers with higher ecological theory scores implemented a wide range of restoration actions and reported relatively fewer positive results but many variable results (Figure 3).
Figure 5. Variation in effectiveness of ecological restoration actions for coastal and inland wetlands in the Great Lakes basin across different monitoring designs. Effects were identified as being variable in 35 papers, positive in 19 papers, or negative (including papers where no effect was observed) in 3 papers (n = 57 papers).

4. Discussion

Published ecological restoration studies in Great Lakes wetland ecosystems increased over time and were most commonly done in the USA. Although a wide range of ecological restoration actions targeting ecological objectives occurred, most studies managed aquatic invasive species with the ecological objective of altering species composition by invasive species removal. These studies integrated ecological theory well within the three stages of restoration despite having vague non-quantitative objectives. The complexity and dynamic nature of restoring and monitoring freshwater ecosystems resulted in the implementation of many study designs using different reference conditions and statistical methods. The effectiveness of ecological restoration actions in relation to ecological objectives in the Great Lakes basin was highly variable. To overcome limitations, explore sources of varying restoration effectiveness, and identify gaps in knowledge, recommendations to improve effective ecological restoration are provided.

4.1. When and Where Has Ecological Restoration Occurred?

Our analysis revealed an increase in published restoration studies over time, that most studies were implemented in the highly degraded lakes Erie, Michigan, and Ontario, and there is a need for large-scale studies to inform effective ecological restoration. Fewer studies in the other Great Lakes (Huron and Superior) reflect the lower level of anthropogenic stressors []. Two thirds of studies occurred in coastal wetlands, likely reflecting their high value, productivity, and functionality despite making up a low proportion of total wetland area in the Great Lakes (e.g., [,]). Almost 60% of studies in the review occurred within individual wetlands ≤ 50 ha in area. Anthropogenic stressors and associated degradation in the Great Lakes basin often occur at larger scales. Small-scale wetland restoration actions may be insufficient when there is a mismatch with the scale of stressors (e.g., []). Mismatch between the restoration actions and stressors within ecosystems was a limitation identified in a review of global freshwater mussel restoration [] and by several studies within our review (e.g., []). In these cases, the authors identified the need to address larger-scale stressors in the face of variable restoration effectiveness. Preliminary studies at proposed ecological restoration sites that assess the relative impacts of small- and large-scale stressors in relation to proposed restoration actions may identify mismatches and ensure the scale of restoration aligns with the scale of relevant stressors.
Peer-reviewed ecological restoration studies were less common on the Canadian side of the basin. Canadian monitoring efforts were only reflected in 20% of the published literature. A global review of ecological restoration found similar results, with limited published research on projects in Canada compared to other countries such as the USA [], and a Canadian review of ecological restoration found that wetlands were less commonly studied compared to other ecosystems such as forests, peatland, grassland, and lakes []. It is possible that the lack of Canadian literature on restoration is due to ~60% of the Great Lakes basin being located within the USA; however, it may also reflect greater financial investments into restoration efforts in the USA compared to Canada, along with greater support for implementing restoration and subsequent monitoring. For example, historical investments by the USA for the restoration of Great Lakes Areas of Concern have been in the billions, compared to millions invested by Canada []. Challenges to the implementation and monitoring of restoration by conservation authorities (one of the primary practitioners in Ontario, Canada) are substantial: reflective of limited resources and organizational mandate (e.g., []). The situation is not unique to Canada, as a recent global review on lake restoration cited lack of support, finances, and weak policy and governance as major reasons for failed restoration [].
Lack of monitoring of restoration actions, when paired with a poor understanding of ecosystem functioning, is a challenge in the broader field of restoration. For example, river restoration efforts in the USA are typically done at small scales, where outcomes are often not monitored []. Limited monitoring efforts inhibit the evaluation of restoration actions in the Great Lakes basin where ecological functions in restored wetland ecosystems are often affected by anthropogenic stressors, the impacts of which are not well understood []. The Great Lakes Coastal Wetland Monitoring Program (2011–2020) is one positive example of bi-national long-term monitoring for ecosystem health, using water, habitat, and biological community indicators for Great Lakes coastal wetlands []. The program can be used as an example for future long-term monitoring efforts. An evaluation of successful restoration practices and funding frameworks in the USA may also provide guidance to address limited monitoring in the Great Lakes basin. To complement long-term monitoring and effectively allocate limited resources, broad-scale analyses should occur to identify candidate sites for restoration actions and monitoring, providing information to aid in the development of lake-wide restoration plans []. When paired, these two recommendations would encourage the scale of restoration actions to expand, aligning better with the scale of stressors and providing broad-scale direction for restoration practitioners [].

4.2. What Approaches Were Taken to Implement and Evaluate Ecological Restoration?

While a wide range of actions were done to address various ecological objectives, the most common action was to manage invasive species with the objective of altering species composition. The upward trend in number of projects undertaking invasive species management over time aligns with the increasing prevalence of aquatic invasive species in the Great Lakes basin [,] and their known negative impacts on a wide range of taxa, ecological communities, and ecosystem functions (e.g., []). The remaining ecological restoration actions also followed an upward trend over time, likely reflecting the broader trend of greater investments into restoration efforts. When examining community-based studies further, ecological theories, commonly succession (e.g., []), resilience (e.g., []), and stable states (e.g., []), were well integrated across all stages of ecological restoration. For studies that did not fully integrate ecological theory, the action phase of studies was often missing theoretical support (e.g., []). At a time when creating sustainable and resilient ecosystems within a socio-ecological landscape (e.g., [,]) has been deemed most effective in practice (e.g., [,]), ecological theory should be tested through field-based experiments where ecological restoration is occurring. The integration of ecological theory into the action phase of restoration allows for the exploration of the complex and dynamic nature of ecosystems. In doing so, the link between theoretical understanding and practice can be strengthened, and the need to ensure that freshwater ecosystems are resilient to future anthropogenic stress can be addressed [].
Despite ecological theory being generally well integrated in the stages of ecological restoration, poorly articulated ecological objectives may explain the lower prevalence of ecological theory in the action stage of restoration and some of the reported variable effectiveness. It is possible that alternative methods of science-based planning and objective setting were used during the action stage but not stated in the studies reviewed. Sixty percent of studies did not contain pre-monitoring data to inform the action stage of ecological restoration, likely explaining, in part, the absence of ecological theory in the action stage. Effective ecological objectives should be founded in ecological theory, quantitative, and clearly linked with practitioner objectives []. Setting quantitative goals for restoration actions often result in positive outcomes [,] whereas the lack of defined, quantitative theory-based ecological objectives can lead to monitoring designs that are not based in scientific evidence and are, as a result, ineffective. Most papers reviewed in this study presented non-quantitative ecological objectives. For example, a common objective was to examine the response of biodiversity or species richness in restored wetlands (e.g., [,]). It is not clear what amount of improvement in biodiversity defines success and what thresholds of increasing biodiversity are meaningful and practical (i.e., []). We expect increases in biodiversity in newly restored wetland ecosystems given their early successional age post-disturbance (e.g., [,]); however, increases do not always reflect better quality habitat or a successful restoration outcome (e.g., [,]). For example, although Phragmites management in coastal wetlands improved species richness and floristic quality, Phragmites re-emergence and the establishment of other invasive plant species resulted in the need for additional action to improve the restored, low-quality plant community []. To inform meaningful biodiversity targets and quantitative ecological objectives, quantitative lake-specific benchmarks that are informed by long-term monitoring programs (e.g., Great Lakes Coastal Wetland Monitoring Program) or modeled using large-scale ecological restoration projects should be developed.
A variety of monitoring designs evaluating the effectiveness of restoration actions in relation to ecological objectives were identified across studies. The BACI study design is known as a robust study design that can separate the recognized natural range of variation across space and time from the ecological changes caused by the restoration actions (e.g., []). To implement a BACI study, pre- and post-project data are required, but are rarely available [,]. Over 70% of studies undertook efficacy monitoring [] that could not account for natural variation (i.e., not BACI studies), and long-term consistent pre- and post-activity monitoring was rare. High proportions of alternative study designs may reflect studies occurring in areas where reference conditions are more difficult to sample (i.e., the rarity of positive reference conditions in Phragmites dense wetlands []) or where pre-restoration data were not available (e.g., []). In some of these cases, study design was strengthened through replication [], paired designs [], or a mix of reference conditions for comparison []. Furthermore, only three of the 16 BACI studies monitored the pre-restoration condition for more than one year, and only five studies monitored the post-restoration response for more than two years. The short-term nature of monitoring may fail to identify long-term ecosystem responses to restoration or when positive effects are reversed after restoration activity stops (e.g., [,,]). Therefore, expanding the temporal and spatial scales of monitoring using robust study designs would provide more confidence in the effectiveness of ecological restoration.
Studies commonly used either a current poor quality and highly impacted reference condition, or a current naturalized and minimally impacted reference condition for comparison. The former approach was particularly common when evaluating invasive species management, where negative reference conditions were used, such as dense monoculture plots of invasive species like Phragmites and hybrid Typha (e.g., []). It is not clear if a positive result in relation to a negative reference is ecologically meaningful because reviews have shown that restoration is effective when compared to negative reference conditions but is not effective when compared to naturalized positive reference conditions []. Furthermore, the classification of positive, minimally impacted reference conditions should be approached with caution given the scale of anthropogenic change in the Great Lakes basin []. Defining appropriate reference conditions (referred to as the reference concept []), has been recognized as a challenge in the broader restoration field, where alternative approaches, including the use of best professional judgement to define a reference community have been implemented []. Proposed approaches also include the use of large-scale, long-term monitoring programs for the development of reference conditions (e.g., Great Lakes Coastal Wetland Monitoring Program) and the development of historical reference conditions from previous records or favorable future reference conditions linked to ecosystem functioning. Additionally, these developed reference conditions could be used to evaluate whether reference conditions in existing studies are appropriate for comparison. Evaluating the accuracy, meaningfulness, and practicality of reference conditions is a current knowledge gap in the restoration field, and no studies in this review formally evaluated these measures.
A wide range of statistical methods were used to evaluate the success of ecological restoration actions in relation to their ecological objectives, and a pattern of increasing complexity was observed over time. The use of many statistical methods may reflect the diversity of study designs used to evaluate a wide range of ecological objectives within the review and/or difficulties encountered when dealing with ecological data (i.e., skewed or zero-inflated data where assumptions cannot be met [], low and/or uneven sample sizes limiting methods and impacting power []). In this review, study designs were often made more robust by increasing replication and/or incorporating multiple references for comparison (e.g., []). However, these robust study designs often include multiple comparisons, increasing the likelihood of obtaining a positive result by chance. This limitation should be overcome by using a correction for multiple comparisons (e.g., [,]). It was also common for studies to use more complex multivariate methods, such as ordination techniques (commonly NMDS (e.g., [])). However, the inclusion of data and model preparation methods were often vague. These choices can affect statistical and ecological conclusions (e.g., [,]). For example, in community-based analyses, the inclusion or exclusion of rare species has been shown to greatly influence site-level results []. Studies should clearly state such choices, indicate whether the assumptions of statistical tests have been met, and evaluate the impacts of common statistical decisions on outcomes (i.e., sensitivity analyses). Lastly, although most studies evaluated community composition, very few considered the impacts of imperfect species detection, where absences are not true absences, but are simply non-detections, and whose consideration can greatly impact the outcomes of a study and subsequent management action (e.g., [,]). Recognizing limited resources, short timelines, and constraints that do not allow for perfect study design, well-invested case studies using model or study ecosystems may serve as surrogates for rigorously testing statistical choices and defining meaningful outcomes.

4.3. How Effective Was Ecological Restoration?

Ecological restoration activities were found to be largely variable in their effectiveness and few patterns emerged as a result. Variable effectiveness was most commonly observed within studies implementing wetland creation (88%), agricultural reversal (67%), or invasive species management (62%). Hydrological management actions had greater success with almost even occurrences of positive (45%) and variable (55%) outcomes. The only action resulting in mostly positive outcomes were studies that implemented multiple restoration actions (56%). Studies that showed negative effectiveness of restoration actions all had objectives that targeted abiotic components of the ecosystem and utilized simple summary or univariate statistics. For example, small-scale restoration actions in two Michigan wetlands did not result in improved water quality [] and larger-scale restoration efforts in the Detroit River, although resulting in some habitat improvement, were not extensive enough to achieve the objective of reaching the required area of soft shoreline []. For studies with a component of community analysis and a low integration of ecological theory, positive restoration effectiveness was almost exclusively observed. For example, restoration projects completed through federal conservation programs in the USA had similar communities in restored and reference wetlands and were successful in providing habitat for Species of Greatest Conservation Need (i.e., []). All observations emphasize the complex nature of ecosystem functions, processes, and health that can be difficult to quantify in simple outcomes. The complex response of an ecosystem to restoration was evident in many studies reporting variable results when monitoring multiple abiotic and biotic responses to the management of invasive species or monitoring responses over time (e.g., Phragmites management []). Novel approaches to monitoring and evaluating ecological restoration may be more sensitive or provide greater clarity to understanding the mechanisms of ecosystem change where variable results are discovered. For example, a taxonomic-diversity approach can be combined with a functional-diversity approach, a method that uses traits related to ecological function (e.g., [,]). In doing so, a better understanding of restoration effectiveness and the mechanisms driving outcomes may be achieved. To better understand ecosystem functioning in relation to ecological objectives, studies targeting the mechanisms of change and tools for evaluating said mechanisms should be developed and tested.
Important knowledge sources outside of the primary literature should also be considered when evaluating ecological restoration across the Great Lakes basin. Restoration and monitoring studies and knowledge are also represented in non-academic sources such as grey literature (e.g., []) and Traditional Ecological Knowledge (TEK). Restoration actions on the Canadian side of the Great Lakes basin are often done by conservation authorities, whose ability to monitor restoration, analyze data, and publish results are often limited by a lack of resources (e.g., []). However, restoration practitioners, such as conservation authorities, possess valuable watershed-based knowledge and insights for undertaking restoration within a socio-ecological landscape. Restoration practitioners have valuable experience and knowledge building relationships with landowners considering restoration (e.g., [,]), often navigating the tension between environmental and private-property protection []. This experience allows them to make informed recommendations for restoration opportunities that align with both ecological and societal priorities []. TEK is another source of knowledge that has largely been unincorporated in ecological restoration, despite the expertise spanning across generations.
To overcome limitations of published research availability for Canadian Great Lakes wetland projects, and to recognize knowledge outside of published literature, collaboration and cooperation must remain central to future restoration efforts (e.g., []). A review of grey literature, followed by structured interviews with existing restoration practitioners, is recommended. Building meaningful relationships with Indigenous communities and providing opportunities for knowledge sharing and co-production are steps that would help address gaps of knowledge in the field of restoration and, most importantly, work towards reconciliation []. TEK has been successfully applied within the Great Lakes basin through invasive species management and commercial fisheries lenses (e.g., [,]). Febria et al. [] also embraced knowledge sharing and co-production through restoration by taking a plural approach, where meaningful relationships between stakeholders and long-term restoration actions were created. Therefore, by considering and exploring alternative sources of knowledge, perspective will be gained that can lead to a more comprehensive approach to restoration. This is key at a time when social and environmental factors must be considered in restoration [] to ensure that restoration is implemented in just and equitable ways.

4.4. Recommendations

Based on the findings of this review and the trends and patterns observed, we provide a set of overall and stage-specific recommendations.

4.4.1. Overall Recommendations

  • Expand the spatial coverage of restoration through increased binational and Canadian efforts for ecological restoration and evaluation;
  • Continue the integration of ecological theory across all phases of ecological restoration (planning, action, and evaluation);
  • Develop lake-specific benchmarks to guide the development of quantitative, meaningful, and practical objectives for ecological restoration; and
  • Expand the ecological restoration knowledge base to include valuable practitioner knowledge and TEK.

4.4.2. Planning Stage of Restoration

  • Invest more substantially in the planning stage of restoration to develop science-based ecological restoration plans and improve study design and statistical analyses (see Figure 6); and
    Figure 6. A decision tree for restoration practitioners and researchers to guide the choice and implementation of study design while recognizing the limitations of what can be studied within each context. A historical or classical approach assumes that ecosystems should be restored to a natural or historical state of equilibrium, often using a known reference or historical condition as a benchmark [,] whereas the modern, novel, or path-dependence approach assumes that ecological functions, processes, and health should be restored within the context of a dynamic socio-ecological landscape, often restoring conditions to those that are resilient to or align with current conditions. Adapted from [].
  • Invest in large-scale and long-term monitoring efforts that can classify existing ecological conditions and ecosystem stressors to identify restoration opportunities.

4.4.3. Action Stage of Restoration

  • Define quantitative, meaningful, and practical ecological objectives for ecological restoration projects; and
  • Continue to invest in BACI study designs that provide insights towards the mechanisms of ecosystem function and change (e.g., functional diversity approach). Alternatively, use well-funded, large-scale ecological restoration projects as model ecosystems to develop effective monitoring and ecological restoration practices. Strengthen small-scale study designs through replication, paired designs, and multiple reference conditions for comparison.

4.4.4. Evaluation Stage of Restoration

  • Monitor ecological restoration over longer time periods (5–10 years at a minimum);
  • Explore novel approaches to defining reference conditions (i.e., developing historical reference conditions based on historical monitoring data and expert knowledge or developing reference conditions that are resilient to current and future anthropogenic stressors) and evaluate if reference conditions are meaningful for comparison; and
  • Report more detailed information on statistical methodologies, include limitations of statistical analyses, and evaluate the implications of statistical choices when monitoring ecological restoration (i.e., imperfect detection modelling, sensitivity analyses).

5. Conclusions

Ecological restoration in coastal and inland wetlands of the Great Lakes varied in effectiveness and included a variety of approaches for implementing and evaluating actions. Restoration research was largely concentrated in the most degraded lakes and increased in frequency over the study period. Insights from this review are limited by the comparatively few number of restoration studies in Canadian wetlands and the absence of TEK and practitioner knowledge (not typically found in published research). To promote successful ecological restoration in the future, restoration requires substantially greater investment into the planning stage of restoration. Success requires increasing long-term monitoring efforts, identifying quantitative and clear restoration objectives that are both meaningful and practical, integrating statistical analyses within project design, and utilizing well-funded restoration opportunities as model ecosystems for field-based experiments. In doing so, we can collaboratively implement science-based restoration approaches, gain a better understanding of the Great Lakes ecosystem, and restore Great Lakes wetlands that are resilient to future anthropogenic stress.

Supplementary Materials

The following supporting information can be downloaded at https://www.mdpi.com/article/10.3390/d17110797/s1, Table S1. Fifty-seven published studies analyzed and classified for a review of ecological restoration actions in Laurentian Great Lakes coastal and inland wetlands. Study types are defined as descriptive (D), before-after (BA), control-impact (CI), or before-after-control-impact (BACI). Ecological theory was classified as either present (1) or absent (0) for each phase of ecological restoration. Refer to the methods for additional information on the classification approach.

Author Contributions

Conceptualization, D.R., S.M.R. and N.E.M.; Methodology, D.R., S.M.R. and N.E.M.; Validation, S.M.R. and N.E.M.; Formal Analysis, D.R.; Investigation, D.R. and S.M.R.; Resources, S.M.R. and N.E.M.; Data Curation, D.R.; Writing—Original Draft Preparation, D.R.; Writing—Review and Editing, S.M.R. and N.E.M.; Visualization, D.R.; Supervision, S.M.R. and N.E.M.; Project Administration, S.M.R. and N.E.M.; Funding Acquisition, S.M.R. and N.E.M. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Canada Nature Fund for Aquatic Species at Risk and NSERC Discovery grant to N.E.M.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

No new data were created or analyzed in this study. Data sharing is not applicable to this article.

Conflicts of Interest

The authors declare no conflicts of interest.

Abbreviations

The following abbreviations are used in this manuscript:
USAUnited States of America
RAPRemedial Action Plan
IUCNInternational Union for Conservation of Nature
DDescriptive
BABefore-after
CIControl-impact
BACIBefore-after-control-impact
TEKTraditional Ecological Knowledge

Appendix A

Table A1. Criteria used to screen papers on an abstract and full-paper level for inclusion in a review of 57 studies evaluating ecological restoration in Great Lakes coastal and inland wetlands.
Table A1. Criteria used to screen papers on an abstract and full-paper level for inclusion in a review of 57 studies evaluating ecological restoration in Great Lakes coastal and inland wetlands.
General Criteria CategoryDescription
Location—Great Lakes
Watershed/Basin		Canada: Studies must occur in Ontario or Quebec in the Great Lakes—St. Lawrence River primary watershed as defined by the Ontario Watershed Boundary.
USA: Studies must occur in Minnesota, Wisconsin, Illinois, Indiana, Michigan, Ohio, Pennsylvania, or New York.
Study system—WetlandsStudies must occur in or measure properties of coastal or inland wetlands, bogs, peatlands, shallow ponds, or fens within the study boundary.
Study variables—MultipleStudies must measure at least one ecosystem function/service or multi-species/community variable, or a combination of abiotic/biotic variables. No single-species studies or studies monitoring waterfowl were included.
Source type—Peer-reviewedPeer-reviewed literature.
Timeline—1800 onNo restrictions on date of publication. Only considers studies evaluating ecosystems from 1800 on (pre-settlement).
Human-mediated improvement
or management papers		Papers considered in this category must analyze the results of human-mediated alterations or management in field studies. Studies only analyzing pre-activity results, studies exclusively using aerial imagery, lab studies, and modeling studies to predict activity outcomes were not included.
Table A2. Integration of ecological knowledge for studies with a component of community analysis for a review of 57 studies evaluating ecological restoration in Great Lakes coastal and inland wetlands. Ecological theories within the context of community analysis can be defined as theories or ideas that explain or describe the mechanisms, processes, and functions of ecosystems. For example, when evaluating community response to restoration, ecological theory-based studies may test theories of succession/community assemblage, competition, alternative stable states, functional traits, and resilience or test the effectiveness of specific mechanisms, like improving connectivity or interspersion, in relation to said theories (see Table 1 in []).
Table A2. Integration of ecological knowledge for studies with a component of community analysis for a review of 57 studies evaluating ecological restoration in Great Lakes coastal and inland wetlands. Ecological theories within the context of community analysis can be defined as theories or ideas that explain or describe the mechanisms, processes, and functions of ecosystems. For example, when evaluating community response to restoration, ecological theory-based studies may test theories of succession/community assemblage, competition, alternative stable states, functional traits, and resilience or test the effectiveness of specific mechanisms, like improving connectivity or interspersion, in relation to said theories (see Table 1 in []).
IUCN–Five Standards of Practice to Guide
Restoration Activities		Remedial
Action Plan-Stages		Evidence of Inclusion in StageExample
Assessment stage: define existing site, landscape,
and reference conditions.		Stage 1:
describe environmental problems and threats in
restoration area and identify any beneficial
use impairments.		Ecological theory is present in the introduction section of the paper as background knowledge. E.g., … projects often increase interspersion by
dredging and scraping openings in an irregular pattern… improving stand structural diversity and local habitat diversity …to improve habitat heterogeneity. []
E.g., However, wetland functions determine not only what organisms will be supported by the system, but also how ecological succession will proceed…whether restored wetlands function in the same way as intact ‘‘natural’’ wetlands is still in question []
Planning and
design stages:
use conditions explored in assessment stage to define restoration objectives and associated activities. 		Stage 2:
identify actions
for remediation.		Ecological theory is present in the objectives, research question, or hypotheses (if stated), where ecological theory or concepts are specifically being tested. The objectives, research questions, or hypotheses are typically located at the end of the introduction and/or in the methods section of the paper. E.g., We hypothesized that if herbicide management releases native plant species from competition, we would see an increase in species richness, diversity, and floristic quality at managed sites. Additionally, because fire removes much of the standing dead biomass that remains after herbicide treatment, increasing light and promoting regeneration from the seedbank, we expected higher richness, diversity, and floristic quality in sites that were burned following herbicide treatment. Alternatively, herbicide management might fail to eradicate Phragmites or promote other invasive species []
E.g., In this paper, we examine the composition of wetland plant assemblages and seed banks in seven diked wetlands along the Great Lakes shoreline and seven nearby wetlands that are still connected to the lake. Our objective is to test the hypothesis that diking significantly affects the vulnerability of coastal wetlands to invasive species []
Monitoring and
evaluation stages:
evaluate the effectiveness
of restoration in relation
to restoration objectives.		Stage 3:
provide evidence that beneficial use impairments have been restored.		Ecological knowledge is present in the discussion to explain observed results. E.g., Lower macroinvertebrate abundance within Typha- dominated control treatments was likely to have been related to structural homogeneity and the associated low abundance of submerged aquatic plants. Although macroinvertebrate density can increase as vegetation structural complexity increases…macroinvertebrates benefit from the highly dissected leaves of many submerged aquatic plants, which increase habitat complexity []
E.g., Plant species richness was more than two times greater at managed sites, indicating that increased light and reduced competition following herbicide treatment promotes germination from the seedbank and/or rapid colonization of many species []
Table A3. Description of reference-condition categories used when classifying studies evaluating ecological restoration in Great Lakes coastal and inland wetlands (n = 57).
Table A3. Description of reference-condition categories used when classifying studies evaluating ecological restoration in Great Lakes coastal and inland wetlands (n = 57).
Type of Reference ConditionDescription of Reference Condition
Historical, pristine reference conditionConditions that are representative of or similar to what the restoration conditions would have been without changes or disturbances caused by anthropogenic activities [], also referred to as pre-degradation reference conditions []. Reference conditions were considered historical when the data was collected > 20 years prior to the implementation of restoration activities. The 20-year timeline aligns with the maximum time frame that the Open Standards for the Practice of Conservation (version 5.0) recommends for achieving conservation goals.
Current, positive, naturalized
reference condition		Conditions that are representative of or similar to what the restoration conditions would have been without changes or disturbance caused by anthropogenic activities [], also referred to as pre-degradation reference conditions []. Reference conditions are considered current when the data was collected in close proximity to the implementation of restoration activities (<20 years).
Current negative, disturbed
reference condition		Pre-restoration conditions [] or conditions that are representative of or similar to restoration conditions and that are in a degraded state due to anthropogenic activities, also referred to as negative reference conditions, post-degradation reference conditions [], or negative controls []
No reference
condition		Some types of studies, by design, do not make formal comparisons to reference conditions and instead focus on analyzing the current state of an ecosystem (i.e., descriptive studies (D)). In these cases, there is no reference condition for formal comparison.

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