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- Wetland restoration ecology - WUR.
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Guided by ecological theory, we develop knowledge on novel restoration measure-effect relations. Examples include building new islands Marker Wadden in lake Marker and rehabilitating stream valley marshes. Furthermore, the effects of climate and global change are taken into account.
Forested riparian zones appeared to be effective to mitigate the stream water temperature rise in small streams induced by climate change in North-western Europe. Furthermore, among other benefits, it helps to maintain or restore the habitat for coldstenothermic stream organisms. We study the effects of shaded-to-open and an open-to-shaded transitions on stream water temperature and on the biological communities inhabiting these streams.
Fine sediments are abundant in urbanized deltas around the world and provide a potential source of building material for wetland construction. In the Netherlands, the MarkerWadden project has embarked aiming at creating a dynamic wetland system in lake Markermeer with gradients in topography, sediments, and rich benthic and wetland biodiversity. Fine sediments will be accumulated into atolls, which eventually develop into valuable ecosystems. We study the interplay of biological, chemical and physical interactions in lake Markermeer.
The parallel assessment of these multiple associations allowed us to capture the simultaneous supply of several ES  , .
- Wetland restoration ecology.
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To avoid counting the same data more than once in a meta-analysis, we performed a separate meta-analysis for each ES using a random-effects model. We considered this approach suitable because we wanted to evaluate each ES separately, rather than the heterogeneity among different ES. We assessed the correlation between biodiversity recovery and ES recovery using the Spearman rank coefficient to quantify the correlation between the corresponding RRs.
We used only RRs from studies that evaluated both biodiversity and ES, and we treated each of these studies as an independent sample. When the same study reported multiple measures of biodiversity or ES, the related RRs were averaged to generate an overall RR for biodiversity and an overall RR for ES for each study, thereby minimizing the risk of pseudo-replication.
This approach led us to combine the four major ES types in order to ensure adequate sample size . We used linear mixed-effects models to evaluate whether the effects of restoration on biodiversity and ES varied with context. Context was parameterized using four nominal fixed factors ecosystem type, main cause of degradation, restoration action, and experimental design and the continuous fixed factor of restoration age, defined as the decimal logarithm of the number of months between completion of the last restoration action and evaluation. We added a fifth nominal fixed factor with two levels biodiversity or ES because we used RRs for both biodiversity and ES recovery in the analysis.
Study site was the random-effect factor and RR was the dependent variable. We also built a second model in which we reduced the degrees of freedom by including only factor categories containing at least 30 observations.
Wetland restoration ecology - WUR
Since this reduced the average sample size in each category, we discarded this model in favor of the first. The selected final model contained main effects but no interactions. All model building and refinement was carried out using Data Desk v6 . The 70 studies analyzed here were distributed across 62 locations in 14 countries Supporting information S4. Restored and natural wetlands showed similar diversity of vascular plants, aquatic invertebrates, macroinvertebrates and protists.
Numbers in parentheses indicate the sample size number of comparisons followed by the numbers of studies. In comparison a , no data were available on non-native vascular plants and protists. In comparison b , the confidence interval for terrestrial invertebrates is not visible because it is smaller than the mean marker. Restoration increased most individual ES that we examined, although not to the same extent Fig. For most individual ES that we examined, restored and natural wetlands tended to supply similar amounts Fig.
Biodiversity and ES response ratios positively correlated in comparisons of restored and degraded wetlands Fig. Comparison of restored and degraded wetlands showed that restoration effects depended on the following factors, listed in order of decreasing importance: main cause of degradation, restoration action, experimental design, and ecosystem type Table 2. In contrast, restoration age did not significantly affect restoration outcomes. These results were the same for the two outcomes of biodiversity recovery and ES recovery.
Context variables explained relatively little variance Restoration significantly ameliorated all causes of degradation that we examined, except for the presence of invasive species Fig. Seven of the 10 restoration actions reported by the included studies showed significant effects on biodiversity and ES supply Fig. Of all restoration actions examined, exotic species removal was associated with the lowest effect size, which did not achieve statistical significance.
Comparison of restored and natural wetlands showed that restoration significantly improved recovery of biodiversity and ES supply Table 2 , although as before, the final model explained only a fraction of the variance Our global meta-analysis, including70 studies conducted in 14 countries, shows that wetland restoration increased biodiversity in degraded wetlands, consistent with another global meta-analysis of different ecosystem types .
In fact, restoration increased the biodiversity of native organisms to levels similar to those in natural wetlands. To be sure, restoration did not improve biodiversity of all organisms uniformly.
Wetland Restoration and Creation: An Overview
Restoration increased vertebrate diversity to levels above those in natural wetlands, though this result may only be transient, since vertebrate richness can vary substantially over time . Conversely, restoration led to levels of biodiversity of non-native vascular plants lower than levels in natural wetlands.
Both of these outcomes may reflect the large, persistent effects of exotic plants on the habitat structure, biodiversity and functioning of wetlands . In addition, wetlands dominated by exotic, invasive plants tend to support fewer native animal species and more invasive animals . Greater diversity by itself is insufficient to ensure high ecosystem functioning . Potentially even more important are the identities and relative proportions of species involved in the restoration process, as well as their ecological and functional properties.
Unfortunately, most studies in our meta-analysis reported aggregate measures of richness or diversity but not community composition Supporting information S1. Indeed a previous meta-analysis of how restoration affects major groups of organisms was restricted to calculating aggregate results for three general categories of vertebrates, macroinvertebrates, and plants .
Higher taxonomic and functional resolution is needed to explore the potentially quite different effects of restoration on organisms that can differ even within a class like vertebrates. Therefore, restoration studies dealing with species composition, community structure and functional ecology are urgently needed. Our meta-analysis showed that restoration enhanced ES supply in degraded wetlands.
The results also showed that it is more difficult to recover ES supply than to recover biodiversity; an alternative or complementary interpretation is that full recovery of ES supply takes longer than full recovery of biodiversity. Either interpretation is consistent with the meta-analysis by Rey Benayas et al. Restoration did not enhance ES uniformly across all individual ES examined. To be sure, we did not expect uniform recovery of all individual ES, given the heterogeneity of ES and wetland types included in the meta-analysis; wetlands types are known to differ in ecological dynamics, recovery rates and extents of recovery .
Our finding that restoration increased supply of provisioning services more than the supply of other ES may reflect the fact that, among the included studies, the desired outcomes when restoring provisioning services e. Effect sizes for these last three services showed wide confidence intervals in our study, suggesting higher intra-class heterogeneity than effect sizes for provisioning services .
Small sample size may explain our finding that restoration did not significantly affect cultural services. Compared to natural wetlands, restored wetlands showed similar supply of provisioning and cultural services but lower supply of regulating services mainly climate regulation, soil fertility and erosion and supporting services mainly biogeochemical cycles and provision of terrestrial habitat.
The lower levels of climate and soil regulation, biological structure and biogeochemical cycles may reflect the intrinsically slow recovery rates reported for these surrogate variables . In contrast, faster recovery rates have been reported for the water regulation variables in our study, such as hydrological dynamics and water quality, and these latter variables indeed showed full recovery.
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Analysis of the ES database, which included abundance data on both non-native plant and animal species, showed that restoration increased regulation of non-native species by reducing their abundance. This result is different than our finding that restoration increased the diversity of such species, though it should be noted that the biodiversity database contained data on non-native plants but not non-native animals.
The abundance of non-native species may decrease rapidly during the restoration process because these species are directly eradicated. However, a reduction in abundance, which reduces the supply of ES, does not necessarily indicate a decrease in species diversity, such as when a habitat contains several rare species in low abundance. Thus, assessment of restoration should take into account both abundance and diversity indicators. The relationship between biodiversity and ES supply remains poorly understood  , yet it is crucial to work out because it has significant implications not only for restoration science but also for wider society, economics, and policy  , .
Our results showed that changes in biodiversity positively correlated with changes in ES supply in a variety of wetlands, ecosystem types and scales, which supports a functional role for biodiversity in the supply of ES  , . This positive relationship is good news for restoration efforts, as it demonstrates the possibility of win-win scenarios for restoring biodiversity and ES. However, such win-win gains have not always proven feasible in practice, especially in restoration projects involving geographically dispersed areas .
Future research should explore how to optimize the synergy between biodiversity and ES supply in the design of management and conservation programs involving restoration. The relationship between biodiversity and ES is also important because it has consequences beyond ecosystem restoration. For example, increasing plant diversity has been shown to enhance the provision of goods from plants and the regulation of erosion, invasive species and pathogens  ; thus, recovering plant diversity may contribute to the recovery of ES beyond the immediate effects of restoration activities.
Future research is needed to disentangle direct and indirect effects of restoration on biodiversity and ES, as well as clarify how the two types of effects interact. Our meta-analysis identified several context factors that significantly affected biodiversity and ES recovery in restored wetlands, including ecosystem type, main cause of degradation, restoration action taken, and experimental design used to assess the restoration. This highlights the need to take context into account when evaluating the effects of wetland restoration. Particularly, examining interaction effects may generate useful insights, but the risk of multiple interactions, including two or even three factors, is too high for the relatively low statistical power of our model.
Our results also showed that biodiversity and ES recovery did not depend on restoration age. Nevertheless, they may depend on how long the restoration process took, on how many times a restoration action was repeated and on the conditions of the degraded wetland prior to restoration. Unfortunately most of the studies included in our meta-analysis did not report such data. The type and duration of interventions required in restoration depend heavily on the type and extent of ecosystem damage .
Future research should examine these context factors in greater detail. Our finding that restoration effects depended on ecosystem type is consistent with an earlier meta-analysis showing that wetlands with more hydrologic flow exchange recovered faster than those that did not receive external water flow .
We obtained different results showing that outcomes of restoration were unrelated to flow exchange, e. Despite these differences, the available evidence strongly indicates that the effectiveness of restoration is habitat-specific, arguing for the need for more research into how to tailor restoration projects to particular environments and how to assess their outcomes accordingly . Our meta-analysis showed that only restoration action determined how close the biodiversity and ES supply of restored wetlands approached those of natural wetlands.
This finding implies that unless the correct restoration action is chosen from the beginning, which is often impossible, the restored wetland may not come as close as possible to natural conditions.
Wetland restoration ecology
Applying a combination of restoration actions may therefore improve the likelihood of success. Taken together, the results of our mixed models suggest that comparisons of degraded, restored, and reference conditions should be carried out to guide and evaluate restoration based on multiple indicators of both biodiversity and ES. These indicators should be consistent with the specific restoration goals  , which can vary greatly depending on the context and project .
Our models further suggest that restoration programs should involve multiple actions to improve the likelihood of success. Comparing degraded, restored and reference conditions to guide restoration may not be feasible in many cases because the irreversibility of much of man-made ecosystem damage makes it difficult to simulate the pre-degradation condition accurately  , and because movement of restored wetlands away from reference conditions makes it difficult to project desired outcomes  , but it should be advisable.
This highlights the need for designing restoration programs with multiple, alternative goals in mind  , . These goals should take into account the social context and human values associated with decisions about wetland management and restoration. The concept of ES can be a robust guide for wetland restoration decision-making because it identifies and quantifies valuable goods and describes the processes and components that provide essential services .
Since several ES are difficult to measure directly, surrogate measures of ecosystem function can be used instead . Accurately assessing the impact of restoration on biodiversity and ES supply requires identifying the particular ecosystem attributes in need of restoration.
To capture potential differences in the restoration of individual ES, we linked the response variables to ES based on specific measures routinely included in ecological studies . In addition, we evaluated the effects of response variables on multiple ES, since the variables may have indirect or unclear links to several ES that significantly affect restoration outcomes. For instance, although all plant species capture carbon, thereby increasing the supply of one ES, non-native species may have detrimental effects on other ES such as biotic interactions.
A restoration action may enhance the supply of one ES while precluding the supply of another  , or it may generate a disservice, such as the release of greenhouse gases. Therefore, analyses of restoration data should assess both the direction and magnitude of associations between response variables and individual ES .
Taking into account the multiple ES associated with a restoration action facilitates the identification of tradeoffs or compromises when planning wetland restoration in which the overriding goal is optimizing multiple ES . Cost plays an important role in restoration planning because it may limit the desired outcomes  , . Surprisingly, the studies included in our meta-analysis did not address the issue of restoration costs.
Costs are an important factor not only during restoration but also after: monitoring of wetlands following their restoration, mitigation or creation is often too brief because it is expensive to evaluate all the ecosystem functions involved. These elements define a complex scenario for decision makers.