Jochem Kail

From Natural to Degraded Rivers and Back Again: A Test of Restoration Ecology Theory and Practice

Extensive degradation of ecosystems, combined with the increasing demands placed on the goods and services they provide, is a major driver of biodiversity loss on a global scale. In particular, the severe degradation of large rivers, their catchments, floodplains and lower estuarine reaches has been ongoing for many centuries, and the consequences are evident across Europe. River restoration is a relatively recent tool that has been brought to bear in attempts to reverse the effects of habitat simplification and ecosystem degradation, with a surge of projects undertaken in the 1990s in Europe and elsewhere, mainly North America. Here, we focus on restoration of the physical properties (e.g. substrate composition, bank and bed structure) of river ecosystems to ascertain what has, and what has not, been learned over the last 20 years. First, we focus on three common types of restoration measures—riparian buffer management, instream mesohabitat enhancement and the removal of weirs and small dams—to provide a structured overview of the literature. We distinguish between abiotic effects of restoration (e.g. increasing habitat diversity) and biological recovery (e.g. responses of algae, macrophytes, macroinvertebrates and fishes). We then addressed four major questions: (i) Which organisms show clear recovery after restoration? (ii) Is there evidence for qualitative linkages between restoration and recovery? (iii) What is the timescale of recovery? and (iv) What are the reasons, if restoration fails? Overall, riparian buffer zones reduced fine sediment entry, and nutrient and pesticide inflows, and positive effects on stream organisms were evident. Buffer width and length were key: 5–30 m width and > 1 km length were most effective. The introduction of large woody debris, boulders and gravel were the most commonly used restoration measures, but the potential positive effects of such local habitat enhancement schemes were often likely to be swamped by larger-scale geomorphological and physico-chemical effects. Studies demonstrating long-term biological recovery due to habitat enhancement were notable by their absence. In contrast, weir removal can have clear beneficial effects, although biological recovery might lag behind for several years, as huge amounts of fine sediment may have accumulated upstream of the former barrier. Three Danish restoration schemes are provided as focal case studies to supplement the literature review and largely supported our findings. While the large-scale re-meandering and re-establishment of water levels at River Skjern resulted in significant recovery of riverine biota, habitat enhancement schemes at smaller-scales in other rivers were largely ineffective and failed to show long-term recovery. The general lack of knowledge derived from integrated, well-designed and long-term restoration schemes is striking, and we present a conceptual framework to help address this problem. The framework was applied to the three restoration types included in our study and highlights recurrent cause–effect chains, that is, commonly observed relationships of restoration measures (cause) and their effects on abiotic and biotic conditions (effect). Such conceptual models can provide useful new tools for devising more effective river restoration, and for identifying avenues for future research in restoration ecology in general.

Influence of large woody debris on the morphology of six central European streams

The impact of large fallen trees on channel form is described for six short stream sections in central Europe influenced by large woody debris (LWD sections), five of which are compared to nearby reference sections free of LWD (reference sections). Three-dimensional models of streambed topography were generated by surveying cross-sections with a spacing of 1 per 1/15 channel width. Parameters derived from digital terrain models and cross-sections compared between LWD sections and reference sections include the extent of pools, bars, and cutbanks, streambed and bank complexity, cross-sectional area, width, depth, and cross-section complexity as described by Andrles [Math. Geol. 26 (1994) 83] ‘angle-measurement-technique’ (AMT analysis), a measure of the deviation of a cross-section line from a straight line. Structural diversity is greater in LWD sections at almost all spatial scales, particularly in terms of pool volume (Mann–Whitney U-test, p<0.01) and cross-section complexity described by median angle of AMT analysis (Mann–Whitney U-test, p<0.05). Large pools are clearly associated with large fallen trees and attain volumes up to 36 m3. With the exception of the ratio of one LWD section where the fallen tree is oriented parallel to flow, the ratio of pool volume to bed planimetric area ranges from 424 to 693 m3/ha, which is in the upper range reported for small, high-gradient streams in Oregon, NW America (229–755 m3/ha) [Can. J. Fish. Aquat. Sci. 47 (1990) 1103]. Pool volume of LWD sections is strongly correlated to the blockage ratio (Spearman rank order correlation, rs=0.93, p<0.01). Differences in channel morphology between the LWD sections and reference sections indicate a strong morphologic control of large woody debris in these central European stream sections.

Contrasting the roles of section length and instream habitat enhancement for river restoration success: a field study of 20 European restoration projects

Restoration of river hydromorphology often has limited detected effects on river biota. One frequently discussed reason is that the restored river length is insufficient to allow populations to develop and give the room for geomorphological processes to occur. We investigated ten pairs of restored river sections of which one was a large project involving a long, intensively restored river section and one represented a smaller restoration effort. The restoration effect was quantified by comparing each restored river section to an upstream non-restored section. We sampled the following response variables: habitat composition in the river and its floodplain, three aquatic organism groups (aquatic macrophytes, benthic invertebrates and fish), two floodplain-inhabiting organism groups (floodplain vegetation, ground beetles), as well as food web composition and land-water interactions reflected by stable isotopes. For each response variable, we compared the difference in dissimilarity of the restored and nearby non-restored section between the larger and the smaller restoration projects. In a second step, we regrouped the pairs and compared restored sections with large changes in substrate composition to those with small changes. When comparing all restored to all non-restored sections, ground beetles were most strongly responding to restoration, followed by fish, floodplain vegetation, benthic invertebrates and aquatic macrophytes. Aquatic habitats and stable isotope signatures responded less strongly. When grouping the restored sections by project size, there was no difference in the response to restoration between the projects targeting long and short river sections with regard to any of the measured response variables except nitrogen isotopic composition. In contrast, when grouping the restored sections by substrate composition, the responses of fish, benthic invertebrates, aquatic macrophytes, floodplain vegetation and nitrogen isotopic composition were greater in sections with larger changes in substrate composition as compared to those with smaller changes. Synthesis and applications. The effects of hydromorphological restoration measures on aquatic and floodplain biota strongly depend on the creation of habitat for aquatic organisms, which were limited or not present prior to restoration. These positive effects on habitats are not necessarily related to the restored river length. Therefore, we recommend a focus on habitat enhancement in river restoration projects. The effects of hydromorphological restoration measures on aquatic and floodplain biota strongly depend on the creation of habitat for aquatic organisms, which were limited or not present prior to restoration. These positive effects on habitats are not necessarily related to the restored river length. Therefore, we recommend a focus on habitat enhancement in river restoration projects.

The importance of the regional species pool, ecological species traits and local habitat conditions for the colonization of restored river reaches by fish

It is commonly assumed that the colonization of restored river reaches by fish depends on the regional species pools; however, quantifications of the relationship between the composition of the regional species pool and restoration outcome are lacking. We analyzed data from 18 German river restoration projects and adjacent river reaches constituting the regional species pools of the restored reaches. We found that the ability of statistical models to describe the fish assemblages established in the restored reaches was greater when these models were based on ‘biotic’ variables relating to the regional species pool and the ecological traits of species rather than on ‘abiotic’ variables relating to the hydromorphological habitat structure of the restored habitats and descriptors of the restoration projects. For species presence in restored reaches, ‘biotic’ variables explained 34% of variability, with the occurrence rate of a species in the regional species pool being the most important variable, while ’abiotic’ variables explained only the negligible amount of 2% of variability. For fish density in restored reaches, about twice the amount of variability was explained by ‘biotic’ (38%) compared to ‘abiotic’ (21%) variables, with species density in the regional species pool being most important. These results indicate that the colonization of restored river reaches by fish is largely determined by the assemblages in the surrounding species pool. Knowledge of species presence and abundance in the regional species pool can be used to estimate the likelihood of fish species becoming established in restored reaches.

Eco-hydrologic model cascades: Simulating land use and climate change impacts on hydrology, hydraulics and habitats for fish and macroinvertebrates.

Climate and land use changes affect the hydro- and biosphere at different spatial scales. These changes alter hydrological processes at the catchment scale, which impact hydrodynamics and habitat conditions for biota at the river reach scale. In order to investigate the impact of large-scale changes on biota, a cascade of models at different scales is required. Using scenario simulations, the impact of climate and land use change can be compared along the model cascade. Such a cascade of consecutively coupled models was applied in this study. Discharge and water quality are predicted with a hydrological model at the catchment scale. The hydraulic flow conditions are predicted by hydrodynamic models. The habitat suitability under these hydraulic and water quality conditions is assessed based on habitat models for fish and macroinvertebrates. This modelling cascade was applied to predict and compare the impacts of climate- and land use changes at different scales to finally assess their effects on fish and macroinvertebrates. Model simulations revealed that magnitude and direction of change differed along the modelling cascade. Whilst the hydrological model predicted a relevant decrease of discharge due to climate change, the hydraulic conditions changed less. Generally, the habitat suitability for fish decreased but this was strongly species-specific and suitability even increased for some species. In contrast to climate change, the effect of land use change on discharge was negligible. However, land use change had a stronger impact on the modelled nitrate concentrations affecting the abundances of macroinvertebrates. The scenario simulations for the two organism groups illustrated that direction and intensity of changes in habitat suitability are highly species-dependent. Thus, a joined model analysis of different organism groups combined with the results of hydrological and hydrodynamic models is recommended to assess the impact of climate and land use changes on river ecosystems.

The role of benthic microhabitats in determining the effects of hydromorphological river restoration on macroinvertebrates

Despite the large number of river restoration projects carried out worldwide, evidence for strong and long-term positive ecological effects of hydromorphological restoration on macroinvertebrates is scarce. To improve the understanding of the success and failure of restoration measures, a standardized field study was carried out in nineteen paired restored and degraded river sections in mid-sized lowland and mountain rivers throughout Europe. We investigated if there were effects of restoration on macroinvertebrate biodiversity, and if these effects could be related to changes in microhabitat composition, diversity and patchiness. Effects were quantified for all taxa combined, as well as Ephemeroptera, Plecoptera and Trichoptera separately. Additionally, species trait classifications of microhabitat preference types were used as a functional indicator. Restoration had no overall positive effects on the selected macroinvertebrate metrics. Rather, we did find positive relationships between the macroinvertebrate responses and the effect of restoration on the diversity and patchiness of microhabitats. Furthermore, the effects on macroinvertebrates could be related to changes in the cover of specific substrate types in the restored sections. We conclude that the limited effect of restoration on macroinvertebrate diversity overall reflected, at least in part, the limited effect of most restoration measures on microhabitat composition and diversity.

Evaluating good-practice cases for river restoration across Europe: context, methodological framework, selected results and recommendations

This introductory paper presents 20 river restoration cases throughout Europe that were investigated in the EU-funded research project REFORM. In the following, this special issue provides seven specific papers that highlight and discuss the effects of restoration on the investigated river–floodplain systems. Additionally, restoration success was estimated from a socio-economic perspective. The first part of this paper presents the overall study concept and the general sampling design of the field investigations. Each study site was examined with the same array of methods, covering habitat composition in the river and its floodplain, three aquatic and two floodplain-related organism groups, as well as food web composition and “aquatic terrestrial” interactions as reflected by stable isotopes. An overview of the rivers and the study sites summarizes main attributes of all investigated sites, with emphasis on the large-scale restoration projects. Some of the projects represent the “state of the art” restoration approaches for two major European river types: gravel-bed mountain rivers and sand-bed lowland rivers. Concluding, restoration efforts had positive effects even in the small restoration projects investigated but did not increase with project size. No “single best” measure could be identified, but river widening generally had a larger effect compared to other restoration measures.

Assessing restoration effects on hydromorphology in European mid-sized rivers by key hydromorphological parameters

The effects of river restoration on hydromorphological conditions and variability are often documented immediately following the restoration, but rarely properly monitored in the long term. This study assesses outcomes of 20 restoration projects undertaken across central and northern Europe for a comprehensive set of hydromorphological parameters, quantified at both larger and smaller spatial scales. For each project, we compared a restored river section to an upstream degraded section. Ten pairs of large projects were contrasted to ten similar but less extensive projects, to address the importance of restoration extent for the success of each project. Overall, river restoration increased habitat diversity through changes in channel morphology. Our results indicated that restoration particularly improved macro- and mesohabitat diversity, but had a limited effect on microhabitat conditions, including the diversity of substrates. We found no significant difference in effects between large and small restoration projects. Our results reveal the need to assess hydromorphological parameters which reflect processes occurring at different spatial scales, including indicators of larger-scale hydromorphological processes such as bank erosion, to monitor restoration effects effectively and accurately. Additionally, our results demonstrate the importance of developing terrestrial parameters, to assess the lateral dimension of river restoration.

FIDIMO — A free and open source GIS based dispersal model for riverine fish

Abstract Dispersal is a key process in community ecology and dynamics that comprise not only various movement processes, but also behavioural components and large-scale spatio-temporal patterns. Modelling fish dispersal further has to consider the species-specific dispersal abilities and the branching, dendritic nature of river networks. With FIDIMO (fish dispersal model) a tool is provided for predicting and simulating spatio-temporal patterns of fish dispersal in river networks with a real integration of GIS for the first time. The model has been written in Python programming language as open source add-on for GRASS GIS. The model has been developed to apply the characteristics of heterogeneous, species and size class specific fish movement on a rasterized river network including migration barriers. Fish dispersal is modelled as a leptokurtic diffusion process spreading from spatially predefined source populations consisting of stationary and mobile components each. The fish dispersal model FIDIMO links conceptual considerations on dispersal modelling with empirically observed fish movement patterns and the strengths of geographically explicit modelling in FOSS GIS. It can be used for different species and any river network meeting the input requirements. As main results FIDIMO yields probabilities of occurrence of fish species in river raster cells based on empirically derived dispersal kernels after the time step of interest modelled. The model output serves to understand and predict time lags and spatio-temporal patterns of recolonization events, the related success of river rehabilitation and also the spread of invasive species. The consequent use of open source programmes allows free access to the code and facilitates easy modifications, adoptions and improvements of the model.

Pressures at larger spatial scales strongly influence the ecological status of heavily modified river water bodies in Germany.

River biota are influenced by anthropogenic pressures that operate at different spatial scales. Understanding which pressures at which spatial scales affect biota is essential to manage and restore degraded rivers. In Europe, many river reaches were designated as Heavily Modified Water Bodies (HMWB) according to the European Water Framework Directive (WFD), where the ecological potential might mainly be determined by pressures at larger spatial scales outside the HMWB (e.g. hydromorphological alterations at the river network and land use at the catchment scale). In Germany, hydromorphological alterations and diffuse pollution were the main pressures. Therefore, the three objectives of this study were to (i) identify the hydromorphological pressures at the site, reach, and river network scale, and land use categories at the catchment scale which significantly affect the ecological status of HMWB in Germany, (ii) quantify the relative importance of these pressures at different spatial scales, and (iii) analyse the differences in response between fish and macroinvertebrates. The results indicated that: (i) At the reach scale, fish were most strongly influenced by channel-bank conditions whilst the naturalness of channel-planform was the best proxy for the ecological status of macroinvertebrates. At the catchment scale, urbanization was the most detrimental land use. (ii) The pressures at larger spatial scales (catchment land use and hydromorphological alterations in the river network) generally were more important than hydromorphological alterations at the reach scale. (iii) Fish were affected equally by both, hydromorphological alterations at the reach scale and large-scale pressures whereas the latter were far more important for the ecological status of macroinvertebrates. In conclusion, these results indicated that large-scale pressures may often limit the efficiency of reach-scale restoration, especially for macroinvertebrates, even in the absence of saprobic pollution, and have to be considered for the management and restoration of HMWB in Germany and comparable degraded river reaches.