This workshop on multivariate analysis of biotic communities focuses on the application of advanced ecological data analysis techniques using R. Participants will gain experience in processing and analyzing data pertaining to biotic communities.

Riverine lakes are a distinct lake type found in geographic regions of Europe shaped by glacial processes. Knowledge concerning their macroinvertebrate communities is scarce and hampers the development of type-specific assessment methods as demanded by the European Union Water Framework Directive (WFD). We compared macroinvertebrate community composition between near-natural riverine lakes and three other types of near-natural lowland lakes to test if communities differ among types. Moreover, we quantified how macroinvertebrate communities from riverine lakes change along a degradation gradient as the basis for developing ecological status classes. Results showed that macroinvertebrate communities significantly differed between riverine and other lowland lake types. The indicator taxa characteristic for near-natural riverine lakes were significantly correlated with environmental characteristics, i.e., comparatively low maximum water depths and high %macrophyte and xylal habitats. Macroinvertebrate communities differed in their taxonomic composition between riverine lake degradation classes, with significant indicator taxa identified for these. We provided empirical evidence that riverine lakes harbour a distinct littoral macroinvertebrate community and recommend establishing riverine lakes as a separate lake type in the Central Plains ecoregion in national typologies for biotic assessments in compliance with the WFD.

Littoral zones − referring to benthic areas above the light compensation depth − provide numerous ecosystem functions, including mediating light, temperature, and nutrient dynamics, and supporting important foraging and refuge areas for macroinvertebrates, fishes and water birds. Habitat assessments of littoral zones remain fundamental to lake and fisheries management, however traditional field surveys are time-intensive and limited in their spatial extent, whereas desktop evaluations using remote sensing and aerial imagery are cost prohibitive and require considerable data processing expertise. In light of these challenges, this study demonstrated the ability to use simple, cost-effective underwater videography to conduct lake-wide spatially-continuous assessments of littoral habitat. For lakes across a gradient of shoreline and riparian development in northwestern United States, we map the areal coverage of macrophytes, coarse woody habitat, bottom substrates, and artificial structures in littoral zones. Underwater videography represents a relevant tool for environmental monitoring because it allows for the estimation of littoral habitats at fine spatial grains across broad spatial extents. Data can also be obtained rapidly and at relatively low cost, providing a permanent record of habitat conditions that can used to monitor trends over time.

Hydromorphological stressors, i.e. hydrological and morphological alterations, affect freshwater biota in running and standing waters in a multitude of ways. These vary depending on size, morphology and other abiotic characteristics of rivers and lakes and also the respective organism groups of which macroinvertebrates, macrophytes and fish are discussed in this chapter. Hydromorphological stressors interact with other stressors, most importantly eutrophication and acidification. Interactions can be characterized as additive, synergistic or antagonistic. Invasive species can modify the effects of hydromorphological alterations. Hydromorphological restorations projects vary considerably in scale, location and effectiveness and more efforts are necessary to design and evaluate these.

The composition of littoral macroinvertebrate communities in lakes is governed by multiple natural and anthropogenic environmental influences interacting at different spatial scales. Since ecological assessment methods using littoral macroinvertebrates should respond specifically to a single stressor, knowledge on the unique effects of a given stressor is necessary. To effectively disentangle the effects of hydromorphology and trophic state requires analysing macroinvertebrate communities at lake sites with the full range of both stressors. We used a dataset of 98 lakes encompassing the entire gradient of geographical locations, lake types, hydromorphological degradation and trophic states in Central European lakes. We studied the unique and joint effects of hydromorphology and trophic state on macroinvertebrate richness, community composition and the Littoral Invertebrate Multimetric Index based on Composite Sampling (LIMCO). Variation partitioning analyses were conducted to test the importance of hydromorphology relative to trophic state across and within hydromorphological states (natural shorelines, hard and soft shore modifications) and trophic states (oligotrophic to hypertrophic states). At natural, hard and soft modification sites, hydromorphology explained 10, 16 and 19%, respectively, of the average unique variation of diversity, community composition and the LIMCO index, whereas trophic state explained on average 2, 5 and 5%, respectively. Similarly, in low, medium and high trophic state lakes, hydromorphology explained 10, 15 and 7%, respectively, of the average unique variation of diversity, community composition and the LIMCO index, whereas trophic state explained on average 0.3, 3 and 6%, respectively. Our results demonstrate that littoral hydromorphology was a more important driver of macroinvertebrate diversity, community composition and LIMCO than trophic state across hydromorphological states and trophic states. This indicates that multiple stressors in lakes act hierarchically on littoral macroinvertebrate communities and that the hydromorphological degradation of littoral zones is the primary driver for altered communities.

The littoral zones of many Central European lakes are severely altered by lake-side retaining walls. These are suspected to impair littoral biota due to the reflection of incoming wave energy. We conducted a comparative study (armoured shore with a retaining wall and a restored shore with a wedge-shaped cobble embankment vs. a pristine shore) at a wind-exposed shore section of Lake Constance. The retaining wall had a number of significant remote effects on the littoral zone, i.e. increased near-bottom current velocities, higher bed-load transport rates, coarsening of surface sediments, reduction in phytomass density, reduction in total densities and number of taxa of macroinvertebrates as well as a significant decline in the percentages of Limnomysis benedeni Czerniavsky, 1882, Ostracoda, Corbicula fluminea O.F. Müller, 1774, Dreissenapolymorpha (Pallas, 1771) and other Bivalvia. However, the significantly affected zone had a rather narrow width of ~ 3 m. Shore restoration measures are needed to remediate negative effects of retaining walls and re-establish ecological conditions that are comparable to those of pristine shores. We recommend a more site sensitive sampling strategy, e.g. for macroinvertebrates in context with the ecological quality assessment under the European Water Framework Directive.

Human lake shore alterations often result in a substantial decrease of littoral and riparian habitat diversity and physical complexity, but the intensity at which shore alterations affect biodiversity may differ among European geographical regions. We tested if the response of littoral macroinvertebrate communities to human shoreline alterations is consistent among geographical regions. We compared community composition and diversity of human altered with those of unmodified littoral zones from 51 lakes across seven European countries in four geographical regions based on pooled composite as well as habitat-specific macroinvertebrate samples. Taxon richness and community composition differed among shore types and different habitats in all geographic regions, with morphological alteration having an overall negative effect on macroinvertebrate taxon richness. In addition, habitat heterogeneity also had a strong effect on littoral communities, with highest taxon richness found in the structurally complex macrophyte habitats in all regions. Average proportional densities of Diptera and Oligochaeta taxa generally increased in morphologically altered shores in all geographical regions, while Bivalvia, Crustacea, Ephemeroptera, Gastropoda and Trichoptera showed comparatively lower numbers in many anthropogenically altered sites. Furthermore, taxon richness was positively correlated with habitat diversity. We were able to relate changes in littoral communities to anthropogenic shoreline alterations, and linked the effect to the loss of habitats and habitat complexity. The results of our study demonstrate that littoral macroinvertebrates respond consistently negative to the influence of morphological alterations across European geographical regions in terms of biodiversity. While macroinvertebrates have previously been identified to be useful descriptors of morphological change in single countries/regions, we can now validate that they can be used to assess the ecological status of lakes in terms of morphological alterations across European regions. Our results can be used to further improve ealready existing WFD-compliant multimetric indices, for example by including taxa groups, which show a strong reaction to shoreline alterations. This could be supported by the inclusion of a suit of indicator taxa reflecting the loss of complex habitats such as macrophytes in the lake littoral.

The spatial distribution of benthic macroinvertebrates along reed transects was studied in lakes with minimal human disturbances to enable a deeper understanding of the functioning of reed macroinvertebrate communities and relations to biotic and abiotic environmental variables. The taxonomic and functional macroinvertebrate community composition significantly differed between outer margin, center, and shore locations. At shore locations, higher proportions of Gastropoda, Hydrachnidia and Coleoptera, mobile swimmers/skaters, predators, and shredders were found. However, outer margin locations were characterized by a higher proportion of sessile filter-feeding Bivalvia and mining Diptera. At the outer margins, also greater contributions of taxa preferring pelal habitats and r-strategists typical for more disturbed environments were observed. An indicator species analysis revealed Asellus aquaticus (Crustacea) and Scirtidae Gen sp. (Coleoptera) as significant indicator taxa for shore locations and Valvata piscinalis (Gastropoda), Tinodes sp. and Orthotrichia sp. (Trichoptera) as significant indicator taxa for outer margin locations. The taxonomic composition of macroinvertebrate communities was significantly related to higher water depth, oxygen content, and pH at outer margin locations. Shore locations were characterized by higher amounts of woody debris, leaf litter, and decaying plant material. In summary, the taxonomic and functional composition of macroinvertebrates varied strongly from shore to outer margin locations and could be related to spatial changes in hydrodynamical and food conditions along the transects.

We examined the effects of woody debris decay state on the densities, taxonomic and functional composition, and diversity of macroinvertebrates in littoral zones of undeveloped lakes. Our study revealed that the interacting effects of increasing roughness and softness of the wood, as well as changes in the quality of food resources with progressing decomposition of woody debris, exerted the strongest impacts on macroinvertebrate communities. Structurally complex, decomposed wood supported higher densities and species richness of macroinvertebrates than undecayed wood. We also found several taxa that were typical for decayed wood, including macroinvertebrates considered as xylophages. Decaying wood underpinned greater functional richness than undecayed wood, with high densities of collector–gatherers and shredders that could benefit from organic matter originating from decomposing wooden tissue, as well as predators attracted by numerous potential prey inhabiting this complex habitat. As decaying wood enhanced abundant and diverse macroinvertebrate communities, which in turn could subsidize upper trophic levels, it provides a valuable habitat in littoral zones, particularly in lakes with already sparse macrophyte cover.

Physical modifications leading to a homogenization of previously diverse littoral habitats increasingly affect the ecological integrity of lake shores in urban landscapes. The European Water Framework Directive (EU WFD) requires integrative assessment of the ecological status of lake ecosystems including lake shore assessment aimed at reaching good ecological status (GES). The ecological consequences of lake shore modifications can be assessed site-specifically by ecological assessment tools based on benthic invertebrates in compliance with the EU WFD. However, it still remains unclear which percentage of the lake shore may be morphologically altered until whole-lake ecological status is affected. We studied a peri-urban lake with similar to 50% of the shoreline altered by urban developments and recreational facilities and the other 50% still remaining in a near-natural, undeveloped state. We assessed the ecological status of each shore type using the Littoral Invertebrate Multimetric Index based on Composite Sampling (LIMCO). Additionally, we used data of a physical habitat survey conducted for each 100-m section within the 12-km long shoreline. We extrapolated site-specific biological assessments to the whole shore length based on pressure-response regressions using physical survey data. Our results showed an overall 'moderate' whole-lake ecological status and consequently the present share of near -natural shoreline is not sufficient to reach GES as required by the EU WFD. GES may be obtained by either further improving existing near -natural shorelines, or by revitalizing developed shorelines. Thus, our approach allows for the quantification of the amount of restoration necessary to derive EU WFD-compliant management objectives for lake shores subjected to human use.

Legislation in Europe has been adopted to determine and improve the ecological integrity of inland and coastal waters. Assessment is based on four biotic groups, including benthic macroinvertebrate communities. For lakes, benthic invertebrates have been recognized as one of the most difficult organism groups to use in ecological assessment, and hitherto their use in ecological assessment has been limited. In this study, we review and intercalibrate 13 benthic invertebrate-based tools across Europe. These assessment tools address different human impacts: acidification (3 methods), eutrophication (3 methods), morphological alterations (2 methods), and a combination of the last two (5 methods). For intercalibration, the methods were grouped into four intercalibration groups, according to the habitat sampled and putative pressure. Boundaries of the 'good ecological status' were compared and harmonized using direct or indirect comparison approaches. To enable indirect comparison of the methods, three common pressure indices and two common biological multimetric indices were developed for larger geographical areas. Additionally, we identified the best-performing methods based on their responsiveness to different human impacts. Based on these experiences, we provide practical recommendations for the development and harmonization of benthic invertebrate assessment methods in lakes and similar habitats.

Lake shores are characterised by a high natural variability, which is increasingly threatened by a multitude of anthropogenic disturbances including morphological alterations to the littoral zone. The European Water Framework Directive (EU WFD) calls for the assessment of lake ecological status by monitoring biological quality elements including benthic macroinvertebrates. To identify cost- and time-efficient sampling strategies for routine lake monitoring, we sampled littoral invertebrates in 32 lakes located in different geographical regions in Europe. We compared the efficiency of two sampling methodologies, defined as habitat-specific and pooled composite sampling protocols. Benthic samples were collected from unmodified and morphologically altered shorelines. Variability within macroinvertebrate communities did not differ significantly between sampling protocols across alteration types, lake types and geographical regions. Community composition showed no significant differences between field composite samples and artificially generated composite samples, and correlation coefficients between macroinvertebrate metrics calculated with both methods and a predefined morphological stressor index were similar. We conclude that proportional composite sampling represents a time- and cost-efficient method for routine lake monitoring as requested under the EU WFD, and may be applied across various European geographical regions.

Morphological degradation constitutes one of the most severe threats to the ecological integrity of lakes. The development of biotic assessment methods for human lake shore alterations using littoral macroinvertebrates requires quantification of the degree of degradation by a stressor index and is complicated through simultaneous physical pressures that alter natural habitat structure.The Lake Habitat Survey (LHS) method and macroinvertebrate sampling were used to produce a pan-European dataset of morphological lake shore degradation and macroinvertebrate densities covering 51 lakes in seven countries and across four geographical regions - northern, western, southern and central Europe.Lake Habitat Survey parameters that differed significantly among three categories of morphological pressure were combined to develop the stressor index components Number of habitats', Habitat diversity', Total percentage volume inhabited by macrophytes', Sum of macrophyte types', Sum of vegetation cover types', Sum of coarse woody debris/roots/overhanging vegetation', Pressure index' (number of human disturbance sources) and Natural/artificial dominant land cover type'.Stressor index components were tested for cross-correlations and for differences among pressure levels. The final composition of the stressor index was optimized for the four studied geographical regions in Europe. The resulting stressor index correlated more strongly with macroinvertebrate metrics than simpler site-specific LHS parameters or the HabQA index developed previously in one lake in north-western Europe.The stressor index developed provides deeper insight into the morphological pressures that affect littoral invertebrate communities. The results also support the use of LHS to quantify morphological stressors at sampling site level, which can ease developing other multimetric bioassessment methods.The stressor index offers the possibility for wide and regional specific application to assess hydromorphological pressures on lakes to assist conservation planning and management and further global efforts to develop and test biotic assessment methods for lakes. Copyright (c) 2014 John Wiley & Sons, Ltd.

Hydromorphological alterations of lake shores constitute an important pressure on Central European lakes. The local ecological effects of such alterations can be assessed by multimetric assessment methods based on eulittoral macroinvertebrate communities. However, such tools so far only enable a local assessment even if the European Union Water Framework (EU WFD) requires an assessment at whole-lake level. Since due to time and financial constraints it is not feasible to perform small-scale macroinvertebrate assessments over the whole length of the lake shore, the site-specific ecological assessment result needs to be extrapolated. In this study, we analyse the use of a physical habitat survey method including both ground surveys and aerial photo analysis (HML = HydroMorphology of Lakes survey protocol) to enable an extrapolation from local to whole lake ecological assessment. For that purpose, we correlated individual macroinvertebrate metrics as well as results of a multimetric macroinvertebrate-based index (I) with the areal cover of surrounding natural shore elements, (II) with the hydromorphological impact within an encircling rectangle around the macroinvertebrate sampling site, and (III) with the hydromorphological impact in the adjacent epi-, eu- and sublittoral subsegments. Building up on this, an ecological whole lake assessment was achieved by averaging multimetric index scores calculated for each subsegement based on a regression equation between the multimetric index and the hydromorphological impacts in the three subsegments. This whole lake assessment approach can most likely be applied to geographical regions outside Central Europe and lake types not analysed here (fluvial lowland lakes, prealpine/alpine lakes), but more studies are necessary to verify this. Eventually, the extrapolation presented here may complement existing lake assessments under the EU WFD solely focusing on water quality so far.

Biomechanical properties and morphological characteristics of stems of eight species of submerged aquatic plants were studied to analyse (1) differences between river and lake specimens, (2) seasonal differences between winter/spring and summer/autumn specimens, and (3) change of biomechanical properties and morphological characteristics along the stems. The data show that river macrophytes display not only characteristic biomechanical traits and morphological characteristics specific to their hydraulic habitats, but also distinctive temporal changes due to seasonally varying water temperature, flow velocity, and growth phase. Furthermore, the data reveal differences between lake and river specimens that could be explained by wind exposure of the lake sampling sites and the species-specific flow requirements of the river macrophytes. Biomechanical properties and morphological characteristics varied along the stem with larger cross-sections and a higher resistance against tension and bending forces at the bottom compared to the top parts, being similar for both lake and river specimens. The acquired and analysed stem biomechanical and morphological data contribute to the plant biomechanics database to underpin a wide range of studies in aquatic ecology, river and wetland management.

The aquatic herbivorous and capital-breeding moth Acentria ephemerella Denis and Schiffermüller, 1775 feeds on submerged pondweeds, Potamogeton spp., and is highly preyed upon by fish in the littoral zone. We studied the spatiotemporal within-lake variability of length, sexual size dimorphism (SSD) and sex ratio of A. ephemerella pupae and of larval population densities. Population densities at three sampling sites strongly increased from July to August and were significantly higher at the Reichenau site in July. Acentria ephemerella sex ratio was male-biased at the Güttingen and Hagnau sites, but showed unbiased or slightly biased sex ratios at Reichenau. The SSD was strongly female-biased. Female size and SSD declined during summer, possibly due to reduced food quantity/quality. The SSD was highest at Reichenau, with little to no differences between Hagnau and Güttingen. At Reichenau, the high population size in July coincided with an unbiased sex ratio, and large SSD/female size due to multiple, possibly interacting factors, including fish predation.

Besides pollution, lakes are affected by human alterations of lake-shore morphology. However, ecological effects of such alterations have rarely been studied systematically. Hence, we developed tools to assess the ecological effects of anthropogenic morphological alterations on European lake-shores based on pressure-specific response patterns of littoral macroinvertebrate community composition. Littoral invertebrates were sampled from 51 lakes in seven European countries. Sampling covered a range of natural to heavily morphologically degraded sites including natural shorelines, recreational beaches, ripraps and retaining walls. Biological data were supplemented by standardized morphological data that were collected via a Lake Habitat Survey (LHS) protocol and subsequently used to develop a morphological stressor index. Two biotic multimetric indices were developed based on habitat-specific samples (Littoral Invertebrate Multimetric based on HAbitat samples, LIMHA) and composite samples (Littoral Invertebrate Multimetric based on COmposite samples, LIMCO) through correlations with the morphological stressor index. Similarity analyses showed strong spatial differences in macroinvertebrate community composition between four main geographical regions, i.e. Western, Northern, Central and Southern Europe. The morphological stressor index as well as LIMCO and LIMHA have been developed for each geographical region specifically, thereby optimizing correlations of LIMCO and LIMHA with the respective morphological stressor index. The metric composition of LIMCO and LIMHA and their correlation coefficients with the morphological stressor index are comparable to existing national and regional methods that assess morphological lakeshore degradation via macroinvertebrate communities. Hence, LIMCO and LIMHA indices constitute a new stressor-specific assessment tool that enables comparable lake morphology assessment across Europe, as it has been developed involving a uniform methodology followed by regionalized optimization. These tools fulfil the standards of the EU Water Framework Directive and thus may complement existing assessment approaches used in lake monitoring focusing solely on lake eutrophication so far. (c) 2013 Elsevier Ltd. All rights reserved.

Implementation of the EU Water Framework Directive (WFD) has drawn much attention to hydromorphological alterations of surface waters. The Lake Habitat Survey (LHS) protocol provides a method for characterising and assessing the physical habitats of lakes and reservoirs. Two metrics were developed based on this method: the Lake Habitat Modification Score (LHMS) and the Lake Habitat Quality Assessment (LHQA), as measures of lake modification and habitat value, respectively. However, the use of these metrics to predict measures of ecological quality remains largely untested. Thus, we assessed the relationships between LHS metrics and the littoral macroinvertebrate community in 42 lakes across Europe. A significant relationship was found between littoral macrophyte descriptors and riparian natural land cover variables of the LHQA score and macroinvertebrate community composition in 2 out of 4 European regions. No relationship was found between macroinvertebrate community composition and the LHMS. Some significant correlations were found between selected macroinvertebrate metrics and the LHS scores, but this pattern was not consistent across regions, and no relationship was found with the overall LHMS or LHQA scores. This demonstrates that the LHS metrics do not consistently predict the quality of littoral macroinvertebrate communities across Europe, and a region specific approach may be necessary. However, we could demonstrate a relationship between the site specific LHS variables and the macroinvertebrate community at the site level, and in some cases at the regional level. Therefore, although the LHS metrics do not appear to be a useful for relating habitat quality and pressure to littoral macroinvertebrate communities, selected LHS variables may exhibit stronger relationships with the biota. (c) 2012 Elsevier Ltd. All rights reserved.

A stereoscopic particle image velocimetry (PIV) system for use in shallow rivers was developed and deployed to study the interactions between turbulent flow and a plant patch in its native environment. Statistical moments of the velocity field were calculated utilising a new method of reducing the contribution of measurement noise, based on the measurement redundancy inherent to the stereoscopic PIV method. The data reveal that the wake of the plant patch was dominated by the presence of a free shear layer induced by the plant drag. Plant motion, estimated from the PlV images, was characterised by travelling waves that propagate along the plant with a velocity nearly equal to the local mean velocity, indicating a direct coupling between turbulence and the plant motion. The characteristic frequency of the plant velocity fluctuations (similar to 1Hz) suggests that the plant motion is dominated by large eddies with dimensions comparable to the flow depth or plant length. Plant and fluid velocity fluctuations were, in contrast, found to be strongly correlated only over a narrow elevation range above the top of the plant, supporting a conjecture that the shear layer turbulence is a key contributor to the plant motion. This talk is devoted to the memory of Professor Stephen E. Coleman - long-term friend and collaborator.