A shift in habitat use patterns of brown trout (Salmo trutta): A behavioural response to macrophyte removal.

Mass development of macrophytes is an increasing problem worldwide and they are frequently removed where they are in conflict with local waterway users. Yet, macrophytes can provide important refuge and nursery habitats for fish. Little is known about the consequences of macrophyte removal for fish behavioural space use and habitat selection. We hypothesised that macrophyte removal would affect brown trout (Salmo trutta) movement during the partial removal of the aquatic plant Juncus bulbosus (L.) in an oligotrophic impounded Norwegian river.We tagged 94 brown trout and tracked them using passive acoustic telemetry for 10 months and mapped the cover of J. bulbosus. Trout behavioural patterns were quantified as space use (utilisation areas 50% and 95%) which was linked to habitat use and selection for J. bulbosus. Removal of J. bulbosus influenced space use of brown trout by reducing the core utilisation area by 22%. Habitat use and selection were likewise influenced by removal with increased use and selection of areas with low J. bulbosus cover (<25%) with corresponding reduction in high J. bulbosus cover (>25-75%). Finally, diurnal differences in space use and habitat use were found, with 19% larger utilisation areas at night and higher use of areas with low J. bulbosus during daytime. Yet, all effect sizes were relatively small compared to the size of the study area. This research provides a detailed case study on the effects of macrophyte removal on fish behavioural patterns in a section of a large Norwegian river with macrophyte mass development. We found no large effects of removal on trout behaviour but noted an increased use of areas with low macrophyte cover. This research is relevant for water managers and policy makers of freshwater conservation and provides a template for using acoustic telemetry to study the effects of macrophyte removal on fish.

Understanding temporal variability across trophic levels and spatial scales in freshwater ecosystems.

A tenet of ecology is that temporal variability in ecological structure and processes tends to decrease with increasing spatial scales (from locales to regions) and levels of biological organization (from populations to communities). However, patterns in temporal variability across trophic levels and the mechanisms that produce them remain poorly understood. Here we analyzed the abundance time series of spatially structured communities (i.e., metacommunities) spanning basal resources to top predators from 355 freshwater sites across three continents. Specifically, we used a hierarchical partitioning method to disentangle the propagation of temporal variability in abundance across spatial scales and trophic levels. We then used structural equation modeling to determine if the strength and direction of relationships between temporal variability, synchrony, biodiversity, and environmental and spatial settings depended on trophic level and spatial scale. We found that temporal variability in abundance decreased from producers to tertiary consumers but did so mainly at the local scale. Species population synchrony within sites increased with trophic level, whereas synchrony among communities decreased. At the local scale, temporal variability in precipitation and species diversity were associated with population variability (linear partial coefficient, ß = 0.23) and population synchrony (ß = -0.39) similarly across trophic levels, respectively. At the regional scale, community synchrony was not related to climatic or spatial predictors, but the strength of relationships between metacommunity variability and community synchrony decreased systematically from top predators (ß = 0.73) to secondary consumers (ß = 0.54), to primary consumers (ß = 0.30) to producers (ß = 0). Our results suggest that mobile predators may often stabilize metacommunities by buffering variability that originates at the base of food webs. This finding illustrates that the trophic structure of metacommunities, which integrates variation in organismal body size and its correlates, should be considered when investigating ecological stability in natural systems. More broadly, our work advances the notion that temporal stability is an emergent property of ecosystems that may be threatened in complex ways by biodiversity loss and habitat fragmentation.

Light and temperature controls of aquatic plant photosynthesis downstream of a hydropower plant and the effect of plant removal.

Many rivers worldwide are regulated, and the altered hydrology can lead to mass development of aquatic plants. Plant invasions are often seen as a nuisance for human activities leading to costly remedial actions with uncertain implications for aquatic biodiversity and ecosystem functioning. Mechanical harvesting is often used to remove aquatic plants and knowledge of plant growth rate could improve management decisions. Here, we used a simple light-temperature theoretical model to make a priori prediction of aquatic plant photosynthesis. These predictions were assessed through an open-channel diel change in O2 mass balance approach. A Michaelis-Menten type model was fitted to observed gross primary production (GPP) standardised at 10 °C using a temperature dependence from thermodynamic theory of enzyme kinetics. The model explained 87 % of the variability in GPP of a submerged aquatic plant (Juncus bulbosus L.) throughout an annual cycle in the River Otra, Norway. The annual net plant production was about 2.4 (1.0-3.8) times the standing biomass of J. bulbosus. This suggests a high continuous mass loss due to hydraulic stress and natural mechanical breakage of stems, as the biomass of J. bulbosus remained relatively constant throughout the year. J. bulbosus was predicted to be resilient to mechanical harvesting with photosynthetic capacity recovered within two years following 50-85 % plant removal. The predicted recovery was confirmed through a field experiment where 72 % of J. bulbosus biomass was mechanically removed. We emphasise the value of using a theoretical approach, like metabolic theory, over statistical models where a posteriori results are not always easy to interpret. Finally, the ability to predict ecosystem resilience of aquatic photosynthesis in response to varying management scenarios offers a valuable tool for estimating aquatic ecosystem services, such as carbon regulation. This tool can benefit the EU Biodiversity Strategy and UN Sustainable Development Goals.

Plastid DNA sequences and oospore characters of some European taxa of Tolypella section Tolypella (Characeae) identify five clusters, including one new cryptic Tolypella taxon from Sardinia, but they do not coincide with current morphological descriptions.

In Europe, the genus Tolypella (Characeae) comprises four to eight Tolypella taxa in sections Rothia and Tolypella that have been distinguished by vegetative morphology and gametangial characters such as antheridial size and oospore wall ornamentation. However, morphological differentiation is difficult in some cases due to overlapping and variable vegetative features, which in many cases are difficult to observe clearly. To clarify the taxonomic status of the five European taxa of Tolypella in section Tolypella, sequence data of the plastid genes atpB, rbcL and psbC for Tolypella glomerata (Desv.) Leonh., Tolypella hispanica Allen, Tolypella nidifica (O.F. Müll.) A. Braun, Tolypella normaniana (Nordst.) Nordst. and Tolypella salina Cor. were combined with data on oospore morphology, including oospore wall ornamentation. Gene sequence data identified five distinct clusters, but they were not consistent with the morphologically identified five taxa. T. glomerata consisted of some of the samples morphologically identified as T. glomerata and seven samples of T. normaniana, while the remaining T. glomerata samples clustered with specimens of unclear affiliation (Tolypella sp.). We identified two clusters of T. hispanica within the European material: cluster T. hispanica I consisted of samples from various locations, whereas the second cluster (T. hispanica II) consisted of samples of T. hispanica from Sardinia Island. The remaining cluster consisted of all the specimens that had been determined as T. salina or T. nidifica in addition to two specimens of T. normaniana. Oospore morphology was most clearly distinguishable for T. glomerata. Oospore characteristics for all other taxa were not as informative but showed some geographical and/or environmentally influenced differences, especially for T. nidifica and T. salina. Our results suggest the need to further check the different taxonomy of Tolypella sect. Tolypella in which specimens normally identified as T. glomerata might be two different taxa, T. glomerata and an unidentified taxon; T. nidifica and T. salina are not separate taxa; T. normaniana is a diminutive variant of two different Tolypella taxa; and T. hispanica comprises two different taxa, one from the Mediterranean island Sardinia.

Short-term effects of macrophyte removal on aquatic biodiversity in rivers and lakes.

Mass development of macrophytes is an increasing problem in many aquatic systems worldwide. Dense mats of macrophytes can negatively affect activities like boating, fishing or hydropower production and one of the management measures often applied is mechanical removal. In this study, we analyzed the effect of mechanical macrophyte removal on phytoplankton, zooplankton, and macroinvertebrate (pelagic and benthic samples) assemblages. Our study covered five sites in four countries in Europe and Africa with highly variable characteristics. In all sites, dense mats of different macrophyte species (Juncus bulbosus in a river in Norway; a mix of native macrophytes in a German river, Elodea nuttallii in a lake in Germany, Ludwigia spp. In a French lake and Pontederia crassipes in a South African lake) are problematic and mechanical removal was applied. In every country, we repeated the same BACI (Before-After-Control-Impact) design, including "before", "one week after", and "six weeks after" sampling in a control and an impact section. Repeating the same experimental design at all sites allowed us to disentangle common effects across all sites from site-specific effects. For each taxonomic group, we analyzed three structural and three functional parameters, which we combined in a scoring system. Overall, the removal of macrophytes negatively affected biodiversity, in particular of zooplankton and macroinvertebrate assemblages. In contrast, plant removal had positive effects on the phytoplankton assemblages. Effects were more pronounced one week after removal than six weeks after. Consequently, we suggest a stronger consideration of the effect of plant removal on biodiversity to arrive at more sustainable management practices in the future.

Relating environmental pressures to littoral biological water quality indicators in Western Balkan lakes: Can we fill the largest gaps?

Along six transects in each of six lakes across the Western Balkans, we collected data for three groups of littoral biological water quality indicators: epilithic diatoms, macrophytes, and benthic invertebrates. We assessed the relationships between them and three environmental pressures: nutrient load (eutrophication), hydro-morphological alteration of the shoreline, and water level variation, separating the effect of individual lakes and continuous explanatory variables. Lake water total phosphorus concentration (TP) showed substantial variation but was not related to any of the tested biological indicators, nor to any of the tested pressures. We suggest that this may be due to feedback processes such as P removal in the lake littoral zone. Instead, we found that a gradient in surrounding land-use towards increasing urbanization, and a land-use-based estimate of P run-off, served as a better descriptor of eutrophication. Overall, eutrophication and water level fluctuation were most important for explaining variation in the assessed indicators, whereas shoreline hydro-morphological alteration was less important. Diatom indicators were most responsive to all three pressures, whereas macrophyte biomass and species number responded only to water level fluctuation. The Trophic Diatom Index for Lakes (TDIL) was negatively related to urbanization and wave exposure. This indicates that it is a suitable indicator for pressures related to urbanization, although a confounding effect of wave exposure is possible. Invertebrate abundance responded strongly to eutrophication, but the indicator based on taxonomic composition (Average Score Per Taxon) did not. Our results suggest that our metrics can be applied in Western Balkan lakes, despite the high number of endemic species present in some of these lakes. We argue that local water management should focus on abating the causes of eutrophication and water level fluctuation, whilst preserving sufficient lengths of undeveloped shoreline to ensure good water quality in the long run.

Mechanical removal of macrophytes in freshwater ecosystems: Implications for ecosystem structure and function.

Macrophytes are generally considered a nuisance when they interfere with human activities. To combat perceived nuisance, macrophytes are removed, and considerable resources are spent every year worldwide on this practice. Macrophyte removal can, however, have severe negative impacts on ecosystem structure and functioning and interfere with management goals of healthy freshwater ecosystems. Here, we reviewed the existing literature on mechanical macrophyte removal and summarised current information from 98 studies on short- and long-term consequences for ecosystem structure and functioning. In general, the majority of studies were conducted in rivers and streams and evaluated short-term effects of removal on single ecosystem properties. Moreover, most studies did not address the interrelationships between ecosystem properties and the underlying mechanisms. Contrasting effects of removal on ecosystem structure and function were found and these discrepancies were highly dependent on the context of each study, making meaningful quantitative comparisons across studies very difficult. We illustrated how a Bayesian network (BN) approach can be used to assess the implications of macrophyte removal on interrelated ecosystem properties across a wide range of environmental conditions. The BN approach could also help engage a conversation with stakeholders on the management of freshwater ecosystems.

Littoral eutrophication indicators are more closely related to nearshore land use than to water nutrient concentrations: A critical evaluation of stressor-response relationships.

Biological assessment metrics and water chemistry measurements are used to quantify the link between stressors and their effects on lake ecosystems, for the Water Framework Directive. However, correlations between metrics and water chemistry are often poor. This is seen as major weaknesses of Water Framework Directive-related monitoring and assessment. We analyzed macrophytes, benthic algae, benthic macroinvertebrates, water chemistry and sediment total phosphorus content in the littoral of six lakes in the Western Balkans and used CORINE land use data to estimate nutrient enrichment via runoff from the adjacent land. Lakes with a higher estimated phosphorus runoff from the adjacent land did not have higher littoral water nutrient concentrations, but littoral diatom assemblages indicated more eutrophic conditions. These lakes also had higher abundances of littoral benthic primary producers, which in turn were associated with low concentrations of dissolved nutrients, but only in autumn, not in spring. This is consistent with primary producers taking up nutrients during the summer growth season. In lakes with high abundances of benthic primary producers, it is likely that the littoral vegetation plays a large role in the transfer of nutrients from the water to the benthos. This process impairs correlations between biological metrics and water nutrient concentrations. Our results suggest that CORINE land cover may be more useful to characterize littoral nutrient enrichment than lake water chemistry. Increased benthic primary producer biomasses and "eutrophic" diatom indices may indicate littoral nutrient enrichment even if water nutrient concentrations are low.

Effects of flow regime on benthic algae and macroinvertebrates - A comparison between regulated and unregulated rivers.

Natural fluctuations in flow are important for maintaining the ecological integrity of riverine ecosystems. However, the flow regime of many rivers has been modified. We assessed the impact of water chemistry, habitat and streamflow characteristics on macroinvertebrates and benthic algae, comparing 20 regulated with 20 unregulated sites. Flow regime, calculated from daily averaged discharge over the five years preceding sampling, was generally more stable at regulated sites, with higher relative discharges in winter, lower relative discharges in spring and smaller differences between upper and lower percentiles. However, no consistent differences in benthic algal or macroinvertebrate structural and functional traits occurred between regulated and unregulated sites. When regulated and unregulated sites were pooled, overall flow regime, calculated as principal components of discharge characteristics over the five years preceding sampling, affected macroinvertebrate species assemblages, but not indices used for ecosystem status assessment or functional feeding groups. This indicates that, while species identity shifted with changing flow regime, the exchanged taxa had similar feeding habits. In contrast to macroinvertebrates, overall flow regime did not affect benthic algae. Our results indicate that overall flow regime affected the species pool of macroinvertebrates from which recolonization after extreme events may occur, but not of benthic algae. When individual components of flow regime were analyzed separately, high June (i.e. three months before sampling) flow maxima were associated with low benthic algal taxon richness, presumably due to scouring. Macroinvertebrate taxon richness decreased with lower relative minimum discharges, presumably due to temporary drying of parts of the riverbed. However, recolonization after such extreme events presumably is fast. Generally, macroinvertebrate and benthic algal assemblages were more closely related to water physico-chemical than to hydrological variables. Our results suggest that macroinvertebrate and benthic algal indices commonly used for ecological status assessment are applicable also in regulated rivers.

DNA barcoding the genus Chara: molecular evidence recovers fewer taxa than the classical morphological approach.

Charophytes (Charales) are benthic algae with a complex morphology. They are vulnerable to ecosystem changes, such as eutrophication, and are red-listed in many countries. Accurate identification of Chara species is critical for understanding their diversity and for documenting changes in species distribution. Species delineation is, however, complicated, because of high phenotypic plasticity. We used barcodes of the ITS2, matK and rbcL regions to test if the distribution of barcode haplotypes among individuals is consistent with species boundaries as they are currently understood. The study included freshly collected and herbarium material of 91 specimens from 10 European countries, Canada and Argentina. Results showed that herbarium specimens are useful as a source of material for genetic analyses for aquatic plants like Chara. rbcL and matK had highest sequence recoverability, but rbcL had a somewhat lower discriminatory power than ITS2 and matK. The tree resulting from the concatenated data matrix grouped the samples into six main groups contrary to a traditional morphological approach that consisted of 14 different taxa. A large unresolved group consisted of C. intermedia, C. hispida, C. horrida, C. baltica, C. polyacantha, C. rudis, C. aculeolata, and C. corfuensis. A second unresolved group consisted of C. virgata and C. strigosa. The taxa within each of the unresolved groups shared identical barcode sequences on the 977 positions of the concatenated data matrix. The morphological differences of taxa within both unresolved groups include the number and length of spine cells, stipulodes, and bract cells. We suggest that these morphological traits have less taxonomic relevance than hitherto assumed.

Interactions between pH and nutrients on benthic algae in streams and consequences for ecological status assessment and species richness patterns.

Eutrophication and acidification are among the major stressors on freshwater ecosystems in northern Europe and North America, but possible consequences of interactions between pH and nutrients on ecological status assessment and species richness patterns have not previously been assessed. Using data from 52 river sites throughout Norway, we investigated the combined effects of pH and nutrients on benthic algae assemblages, specifically 1) taxa-specific couplings between nutrient and acidity traits, 2) the degree of consistency between different biotic indices, separately for nutrients and acid conditions, 3) the impact of pH on nutrient indices and phosphorus on indices of acid conditions, and 4) the impact of pH and phosphorus supply on diatom and non-diatom taxon richness. We found that 1) acid-tolerant taxa are generally associated with nutrient-poor conditions, with only a few exceptions; this is probably more a consequence of habitat availability than reflecting true ecological niches; 2) correlation coefficients between nutrient indices and TP, as well as acid conditions indices and pH were barely affected when the confounding factor was removed; 3) the association of acid-tolerant taxa with nutrient-poor conditions means that the lowest possible nutrient index at a site, as indicated by benthic algae, is lower at acid than at circumneutral sites. Although this may be an artifact of the datasets from which taxa-specific indicator values were derived, it could lead to a drift in nutrient indices with recovery from acidification; 4) the response of non-diatom taxon richness follows a complex pattern with a synergistic interaction between nutrient supply and pH. In contrast, diatom richness follows a simple additive pattern; this suggests structural differences between diatoms and non-diatom benthic algae in their response to nutrient supply and pH; diatom taxon richness tended to increase with nutrient supply, while non-diatom richness decreased.

Bioconcentration and intracellular storage of hexachlorobenzene in charophytes and their potential role in monitoring and remediation actions.

It has been hypothesized that highly hydrophobic substances (LogK(OW) > 5) including many persistent organic pollutants cannot overtake protective tissues and diffuse inside the body of plants due to steric hindrance or very slow diffusion. We investigated the bioaccumulation of hexachlorobenzene (HCB, LogK(OW) = 5.5) in a benthic charophycean macro-alga: Chara rudis. Chara species are a group of common freshwater algae with a complex body structure encompassing a protective layer of cortex cells surrounding large internode cells. The charophyte cell wall has many features in common with that of higher plants; therefore, they are useful models to investigate bioaccumulation mechanisms in general. We found that HCB diffused through the cortex and reached the cytoplam of internode cells. More than 90% of the HCB mass found in the organism was in the cortex and 10% in the internode cell cytoplasm. The cortex reached a pseudoequilibrium partitioning with water, and the bioconcentration factor was in the same range as that of lower aquatic organisms such as phytoplankton. Charophytes are therefore very efficient accumulators of hydrophobic compounds. Based on the structural and ecological features of charophytes, we speculated on their possible use as biomonitors and bioremediation tools.

Impact of calcium and TOC on biological acidification assessment in Norwegian rivers.

Acidification continues to be a major impact in freshwaters of northern Europe, and the biotic response to chemical recovery from acidification is often not a straightforward process. The focus on biological recovery is relevant within the context of the EU Water Framework Directive, where a biological monitoring system is needed that detects differences in fauna and flora compared to undisturbed reference conditions. In order to verify true reference sites for biological analyses, expected river pH is modeled based on Ca and TOC, and 94% of variability in pH at reference sites is explained by Ca alone, while 98% is explained by a combination of Ca and TOC. Based on 59 samples from 28 reference sites, compared to 547 samples from 285 non-reference sites, the impact of calcium and total organic carbon (TOC) on benthic algae species composition, expressed as acidification index periphyton (AIP), is analyzed. Rivers with a high Ca concentration have a naturally higher AIP, and TOC affects reference AIP only at low Ca concentrations. Four biological river types are needed for assessment of river acidification in Norway based on benthic algae: very calcium-poor, humic rivers (Ca<1 mg/l and TOC>2 mg/l); very calcium-poor, clear rivers (Ca<1 mg/l and TOC<2 mg/l); calcium-poor rivers (Ca between 1 and 4 mg/l); moderately calcium rich rivers (Ca>4 mg/l). A biological assessment system for river acidification in Norway based on benthic algae is presented, following the demands of the Water Framework Directive.