Escaping malnutrition by shifting habitats: A driver of three-spined stickleback invasion in Lake Constance.

Fatty acids, and especially long-chain polyunsaturated fatty acids, are biologically important components in the metabolism of vertebrates, including fish. Essential fatty acids (EFA) are those that in a given animal cannot be synthesized or modified from precursors and must therefore be acquired via the diet. Because EFAs are often unevenly distributed in nature, this requirement may drive species to make behavioral or ecological adaptations to avoid malnutrition. This is especially true for fish like the three-spined stickleback (Gasterosteus aculeatus L.) of Upper Lake Constance (ULC), whose recent marine ancestors evolved with access to EFA-rich prey, but which found themselves in an EFA-deficient habitat. An unexpected and unprecedented ecological shift in the ULC stickleback population from the littoral to pelagic zones in 2012 might be linked to EFA availability, triggering ecological release and enabling them to build a hyperabundant population while displacing the former keystone species, the pelagic whitefish Coregonus wartmanni. To test this hypothesis, sticklebacks from the littoral and pelagic zones of ULC were sampled seasonally in two consecutive years, and their stomach contents and fatty acid profiles were analysed. Pelagic sticklebacks were found to possess significantly higher values of an important EFA, docosahexaenoic acid (DHA), especially during autumn. Evaluation of the DHA supply suggests that sticklebacks feeding in the littoral zone during autumn could not meet their DHA requirement, whereas DHA availability in the pelagic zone was surplus to demand. During autumn, pelagic sticklebacks consumed large amounts of DHA-rich prey, that is, copepods, whereas littoral sticklebacks relied mainly mostly on cladocerans, which provide much lower quantities of DHA. Access to pelagic zooplankton in 2012 was possibly facilitated by low densities of previously dominant zooplanktivorous whitefish. The present study offers a convincing physiological explanation for the observed expansion of invasive sticklebacks from the littoral to the pelagic zones of Lake Constance, contributing to a phase shift with severe consequences for fisheries.

Genome-Wide Reconstruction of Rediploidization Following Autopolyploidization across One Hundred Million Years of Salmonid Evolution.

The long-term evolutionary impacts of whole-genome duplication (WGD) are strongly influenced by the ensuing rediploidization process. Following autopolyploidization, rediploidization involves a transition from tetraploid to diploid meiotic pairing, allowing duplicated genes (ohnologs) to diverge genetically and functionally. Our understanding of autopolyploid rediploidization has been informed by a WGD event ancestral to salmonid fishes, where large genomic regions are characterized by temporally delayed rediploidization, allowing lineage-specific ohnolog sequence divergence in the major salmonid clades. Here, we investigate the long-term outcomes of autopolyploid rediploidization at genome-wide resolution, exploiting a recent "explosion" of salmonid genome assemblies, including a new genome sequence for the huchen (Hucho hucho). We developed a genome alignment approach to capture duplicated regions across multiple species, allowing us to create 121,864 phylogenetic trees describing genome-wide ohnolog divergence across salmonid evolution. Using molecular clock analysis, we show that 61% of the ancestral salmonid genome experienced an initial "wave" of rediploidization in the late Cretaceous (85-106 Ma). This was followed by a period of relative genomic stasis lasting 17-39 My, where much of the genome remained tetraploid. A second rediploidization wave began in the early Eocene and proceeded alongside species diversification, generating predictable patterns of lineage-specific ohnolog divergence, scaling in complexity with the number of speciation events. Using gene set enrichment, gene expression, and codon-based selection analyses, we provide insights into potential functional outcomes of delayed rediploidization. This study enhances our understanding of delayed autopolyploid rediploidization and has broad implications for future studies of WGD events.

Freshwater mussel conservation: A global horizon scan of emerging threats and opportunities.

We identified 14 emerging and poorly understood threats and opportunities for addressing the global conservation of freshwater mussels over the next decade. A panel of 17 researchers and stakeholders from six continents submitted a total of 56 topics that were ranked and prioritized using a consensus-building Delphi technique. Our 14 priority topics fell into five broad themes (autecology, population dynamics, global stressors, global diversity, and ecosystem services) and included understanding diets throughout mussel life history; identifying the drivers of population declines; defining metrics for quantifying mussel health; assessing the role of predators, parasites, and disease; informed guidance on the risks and opportunities for captive breeding and translocations; the loss of mussel-fish co-evolutionary relationships; assessing the effects of increasing surface water changes; understanding the effects of sand and aggregate mining; understanding the effects of drug pollution and other emerging contaminants such as nanomaterials; appreciating the threats and opportunities arising from river restoration; conserving understudied hotspots by building local capacity through the principles of decolonization; identifying appropriate taxonomic units for conservation; improved quantification of the ecosystem services provided by mussels; and understanding how many mussels are enough to provide these services. Solutions for addressing the topics ranged from ecological studies to technological advances and socio-political engagement. Prioritization of our topics can help to drive a proactive approach to the conservation of this declining group which provides a multitude of important ecosystem services.

A roadmap for the conservation of freshwater mussels in Europe.

Europe has a long history of human pressure on freshwater ecosystems. As pressure continues to grow and new threats emerge, there is an urgent need for conservation of freshwater biodiversity and its ecosystem services. However, whilst some taxonomic groups, mainly vertebrates, have received a disproportionate amount of attention and funds, other groups remain largely off the public and scientific radar. Freshwater mussels (Bivalvia, Unionida) are an alarming example of this conservation bias and here we point out six conceptual areas that need immediate and long-term attention: knowledge, threats, socioeconomics, conservation, governance and education. The proposed roadmap aims to advance research, policy and education by identifying the most pressing priorities for the short- and long-term conservation of freshwater mussels across Europe.

A global agenda for advancing freshwater biodiversity research.

Global freshwater biodiversity is declining dramatically, and meeting the challenges of this crisis requires bold goals and the mobilisation of substantial resources. While the reasons are varied, investments in both research and conservation of freshwater biodiversity lag far behind those in the terrestrial and marine realms. Inspired by a global consultation, we identify 15 pressing priority needs, grouped into five research areas, in an effort to support informed stewardship of freshwater biodiversity. The proposed agenda aims to advance freshwater biodiversity research globally as a critical step in improving coordinated actions towards its sustainable management and conservation.

Nutrient and fine sediment loading from fish pond drainage to pearl mussel streams - Management implications for highly valuable stream ecosystems.

Man-made, drainable aquaculture ponds have the potential to affect the water quality in the receiving waters, but whether they act mainly as a source or sink of fine sediments and nutrients is still unclear. Particularly in oligotrophic streams containing populations of the highly endangered freshwater pearl mussel (Margaritifera margaritifera), even low additional inputs pose the threat of exceeding thresholds for downstream habitat quality. In this study, the effluent quality during the drainage of two extensively used cyprinid ponds with a size of 0.103 and 0.150 ha was monitored at a high temporal resolution, to characterize the nutrient and sediment loading into the receiving stream under two different management scenarios. The loading of total suspended solids (TSS) was disproportionally dominated by the final step of pond drainage during the fish harvest, when a proportion of 30% of the particles released over the entire drainage process was released with only 1% of the total water volume drained. The continuous release of the ponds' surface water resulted in an additional loading of 28.8 kg/ha of NO3-N, 0.82 kg/ha of NH4-N and 0.58 kg/ha of total-P that was not strongly enhanced by the fish harvest. Using a settling pond was an efficient measure to reduce the amount of suspended particles and excess ammonium and phosphorous reaching the receiving stream. Without such a measure, TSS concentrations in the receiving stream during the fish harvest were elevated to a maximum of >900 mg/l, representing a 20-fold increase compared to 45 mg/l upstream. However, about 1/3 of the released TSS were retained in the overgrown outflow ditch. The differences in loading and retention patterns of dissolved and particulate pollutants revealed the need for divergent approaches to address suspended or dissolved pollutants: Physical settling structures can be effective at reducing particulate inputs, but they might not be sufficient to mitigate the negative effects on oligotrophic streams without a specific design to sustainably remove nutrients. This information on drainage management is not only relevant for minimizing the impacts of aquaculture ponds on downstream ecosystems, but also for the maintenance of nature conservation and flood retention ponds.

The role of anthropogenic habitats in freshwater mussel conservation.

Anthropogenic freshwater habitats may provide undervalued prospects for long-term conservation as part of species conservation planning. This fundamental, but overlooked, issue requires attention considering the pace that humans have been altering natural freshwater ecosystems and the accelerated levels of biodiversity decline in recent decades. We compiled 709 records of freshwater mussels (Bivalvia, Unionida) inhabiting a broad variety of anthropogenic habitat types (from small ponds to large reservoirs and canals) and reviewed their importance as refuges for this faunal group. Most records came from Europe and North America, with a clear dominance of canals and reservoirs. The dataset covered 228 species, including 34 threatened species on the IUCN Red List. We discuss the conservation importance and provide guidance on how these anthropogenic habitats could be managed to provide optimal conservation value to freshwater mussels. This review also shows that some of these habitats may function as ecological traps owing to conflicting management practices or because they act as a sink for some populations. Therefore, anthropogenic habitats should not be seen as a panacea to resolve conservation problems. More information is necessary to better understand the trade-offs between human use and the conservation of freshwater mussels (and other biota) within anthropogenic habitats, given the low number of quantitative studies and the strong biogeographic knowledge bias that persists.

Algal Community Change in Mountain Lakes of the Alps Reveals Effects of Climate Warming and Shifting Treelines1.

The biological communities of mountain lakes are suspected to be highly sensitive to global warming and associated catchment changes. To identify the parameters determining algal community responses, subfossil pigments from 21 different mountain lakes in the Bavarian-Tyrolean Limestone Alps were investigated. Sediment cores were radio-isotopically dated, and their pigment preservation evaluated. General additive models (GAM) of pigment compositions were calculated with temperature as the explanatory variable and generalized linear models with several lake parameters explaining log-transformed GAM P-values. Lake depth and trophic state were identified as major control variables of the algal community and productivity changes. Shifts in a deep oligotrophic alpine lake (lg(P) = -1.04) were half as strong as in a shallow mesotrophic alpine lake (lg(P) = -1.86) with faster warming and higher productivity forcing the development of biomass. Phytoplankton and macrophyte pigments increased clearly with warming, at lower altitudes, and decreased at the treeline, so that periphytic pigments dominated alpine sediments. This pattern is probably the result of interactions of UV radiation and allochthonous inputs of DOM. Our findings suggest that (sub)alpine shallow lakes with higher nutrient levels are most vulnerable to climate change-driven changes whereas deep, nutrient-poor lakes appear more resilient.

Host (Salmo trutta) age influences resistance to infestation by freshwater pearl mussel (Margaritifera margaritifera) glochidia.

The freshwater pearl mussel (Margaritifera margaritifera) is an endangered bivalve with an obligate parasitic stage on salmonids. Host suitability studies have shown that glochidial growth and load vary significantly between host strains as well as among individuals of a suitable strain. Variation in host suitability has been linked to environmental conditions, host age and/or size, genetic composition of the host and parasite, or a combination of these factors. In our study, we wanted to investigate if brown trout (Salmo trutta) displayed an age-dependent response to glochidial infestation. We hypothesised that 1+ naive brown trout hosts tolerate glochidial infestation better than 0+ hosts. In order to test our hypothesis, we infested 0+ and 1+ hatchery reared brown trout with glochidia from closely related mothers and kept them under common garden conditions. This allowed us to observe a pure age dependent host response to infestation, as we eliminated the confounding effect of genotype-specific host interactions. We analysed the interaction between glochidial load and host condition, weight and length, and observed a significant age-dependent relationship. Glochidial load was negatively correlated to host condition in 0+ fish hosts and positively correlated in 1+ hosts. These contradictory findings can be explained by a change in host response strategy, from resistance in young to a higher tolerance in older fish. In addition, we also examined the relationship between glochidial load and haematocrit values in the 1+ hosts and observed that haematocrit values were significantly higher in heavily infested hosts. Our results have important conservation implications for the management of wild pearl mussel populations, as well as for captive breeding programmes.

Boom-bust dynamics in biological invasions: towards an improved application of the concept.

Boom-bust dynamics - the rise of a population to outbreak levels, followed by a dramatic decline - have been associated with biological invasions and offered as a reason not to manage troublesome invaders. However, boom-bust dynamics rarely have been critically defined, analyzed, or interpreted. Here, we define boom-bust dynamics and provide specific suggestions for improving the application of the boom-bust concept. Boom-bust dynamics can arise from many causes, some closely associated with invasions, but others occurring across a wide range of ecological settings, especially when environmental conditions are changing rapidly. As a result, it is difficult to infer cause or predict future trajectories merely by observing the dynamic. We use tests with simulated data to show that a common metric for detecting and describing boom-bust dynamics, decline from an observed peak to a subsequent trough, tends to severely overestimate the frequency and severity of busts, and should be used cautiously if at all. We review and test other metrics that are better suited to describe boom-bust dynamics. Understanding the frequency and importance of boom-bust dynamics requires empirical studies of large, representative, long-term data sets that use clear definitions of boom-bust, appropriate analytical methods, and careful interpretations.

Synechococcus diversity along a trophic gradient in the Osterseen Lake District, Bavaria.

Picocyanobacteria are important primary producers in freshwater; however, there is still a knowledge gap regarding their diversity at the strain level. For this reason, the microbial diversity of four lakes with different trophic states was investigated by sequencing of the 16S rRNA gene using universal primers. The study was performed in selected lakes of the Osterseen Lake District, Germany, from 2012 to 2014 (Lake Schiffhuettensee: eutrophic; Lake Ostersee: meso-oligotrophic; Lake Groebensee: oligotrophic; Lake Lustsee: oligotrophic). It was determined that the bacterial community of each of these lakes was characterized by one or more specific phyla. Within the autotrophic plankton, the picocyanobacterium Synechococcus sp. dominated oligotrophic habitats, whereas eukaryotic algae prevailed in eutrophic lakes. The study focused on the occurrence of cyanobacteria, specifically the genus Synechococcus. Genetic analysis of the 16S rRNA gene revealed an extendend diversity of freshwater Synechococcus. The occurrence of the identified operational taxonomic units of Synechococcus did not correlate with the trophic state of their habitat, suggesting that the current, underestimated diversity of picocyanobacteria deserves increased consideration in assessments of microbial and freshwater biodiversity.

The use of growth and behavioral endpoints to assess the effects of pesticide mixtures upon aquatic organisms.

Aquatic communities are often subject to complex contaminant mixtures, usually at sublethal concentrations, that can cause long-term detrimental effects. Chemicals within mixtures can effectively interact, resulting in synergism, antagonism or additivity. We investigated the tertiary mixture effects of two pyrethroids, lambda-cyhalothrin and permethrin, and the organophosphate chlorpyrifos, evaluating sublethal endpoints; immobility and growth, on Chironomus dilutus in 10-day exposures. We utilized a toxic units (TU) approach, based on median lethal concentrations (LC50) for each compound. The concepts of independent action and concentration addition were used to compare predicted mixture toxicity to observed mixture toxicity. Increased immobility resulted from mixture concentrations ≥1 TU (7.45 ng/L lambda-cyhalothrin × 24.90 ng/L permethrin × 129.70 ng/L chlorpyrifos), and single pesticides concentrations ≥0.25 TU (5.50 ng/L lambda-cyhalothrin, 24.23 ng/L permethrin, 90.92 ng/L chlorpyrifos, respectively). Growth was inhibited by pesticide mixtures ≥0.125 TU (1.04 ng/L lambda-cyhalothrin × 3.15 ng/L permethrin × 15.47 ng/L chlorpyrifos), and singly by lambda-cyhalothrin ≥0.25 TU (5.50 ng/L), and permethrin ≥0.167 TU (18.21 ng/L). The no observed effect concentrations (NOEC) for immobility and growth, for both mixture and single-pyrethroid exposure, were up to 8.0 and 12.0 times respectively lower than the corresponding NOEC for survival. The median effective concentrations (EC50) for growth (mixture and single-pyrethroid exposure) were up to 7.0 times lower than the respective LC50. This study reinforces that the integration of sublethal endpoints in monitoring efforts is powerful in discerning toxic effects that would otherwise be missed by solely utilizing traditional toxicity assessments.

Characterization of the reproductive strategy of invasive Round Goby (Neogobius melanostomus) in the Upper Danube River.

Originating from the Black and Caspian seas, the Round Goby (Neogobius melanostomus) has become one of the most successful invaders of freshwater ecosystems. In this study, we provide a characterization of the reproductive strategy of an established population of Round Gobies in the Upper Danube river including sex ratio, fluctuations of gonadosomatic index (GSI), analysis of timing of spawning as well as of clutch and egg size. We compare these results to other studies from the native and invaded range. In the Danube, the Round Goby population was found to be female dominated, however fluctuations in magnitude of female bias were observed between months. Monitoring of the population across 1.5 years revealed that GSI was highest from April to June, while lowest values were observed in August and September. Using time-series analysis, a delayed effect of temperature on GSI was found for females and males, while a quicker response of GSI levels to photoperiod and discharge was observed for females. GSI increased with body size for females and eggs were found to be significantly larger in May, however clutch sizes did not differ between months. Results of a literature review revealed great differences in timing and length of spawning season as well as sex ratio between populations throughout the distribution range, which can probably be explained by climatic and photoperiodic conditions together with the time since invasion and the high plasticity of Round Gobies.

Multiple climate change stressors reduce the emergence success of gravel-spawning fish species and alter temporal emergence patterns.

Climate change, with its profound effects on stream sediment, hydrological, and temperature dynamics, will exacerbate impacts on habitat conditions for many species, particularly those with vulnerable early life stages relying on the hyporheic zone, such as gravel-spawning fishes. Due to the complex and interactive nature of multiple stressor effects, we employed large-scale outdoor mesocosms to systemically test how the reproductive success of three gravel-spawning fish species brown trout (Salmo trutta), nase, (Chrondrostoma nasus) and Danube salmon (Hucho hucho) was affected by individual and combined effects of warming (+3-4 °C), fine sediment (increase in <0.85 mm by 22 %) and low-flow (eightfold discharge-reduction). Fine sediment had the most detrimental effect on emergence rate and fry length in all three species, reducing the emergence rate to zero in brown trout, 9 % in nase, and 4 % in Danube salmon. The emergence mortality caused by fine sediment surpassed that of hatching distinctly, suggesting that negative effects due to hypoxia were considerably exacerbated by entombment. Warming had only minor effects as a single stressor, but low flow reduced emergence rates of the spring spawning species nase and Danube salmon by 8 and 50 %, respectively. In combined treatments including fine sediment, however, the emergence success of all three species responded strongly negatively, even in the cyprinid species nase, which showed little interactive effects between stressors regarding hatching success. Warming and fine sediment also led to the earlier emergence of fry, implying a risk of asynchrony with available food resources. This study dramatically shows that climate change can have deleterious impacts on the reproductive success of gravel-spawning fish species, irrespective of taxonomic or ecological traits.

Ecosystem services potential is declining across European capital metropolitan areas.

Ecosystem services (ES) are essential to sustainable development at multiple spatial scales. Monitoring ES potential (ESP) at the metropolitan level is imperative to sustainable cities. We developed a procedure for long-term monitoring of metropolitan ESP dynamics, utilizing open-source land use land cover (LULC) data and the expert matrix method. We compared the ESP results of 38 European Capital Metropolitan Areas (ECMA) regarding biodiversity integrity, drinking water provision, flood protection, air quality, water purification, and recreation & tourism. Our results show significant declines in ESP across ECMA due to LULC alteration between 2006, 2012, and 2018. We found that ECMA in post-socialist European countries like Poland (Warszawa) have experienced high rates of land use transformation with a remarkable impact on ESP. Surprisingly, we found that Fennoscandinan ECMA, like Helsinki, Stockholm, and Oslo which lead the cumulative ESP ranking, faced the ESP reduction of the highest impact in recent years. The correlation analysis of ESP dynamics to urban expansion and population growth rates suggests that inattentive urbanization processes impact ESP more than population growth. We unveil the implications of our results to the EU and global level agendas like the European Nature Conservation Law and the Sustainable Development Goals.

Surviving global change: a review of the impacts of drought and dewatering on freshwater mussels.

The increase in the frequency and intensity of droughts and heatwaves caused by climate change poses a major threat to biodiversity. In aquatic systems, sedentary species such as freshwater mussels are generally considered more vulnerable to changes in habitat conditions than mobile species such as fish. As mussels provide important ecosystem services, understanding the impacts of drought on freshwater mussels is of particular importance for the management of overall functioning of aquatic ecosystems. We used a comprehensive literature search to provide a systematic overview of direct and indirect effects of drought on freshwater mussels (Bivalvia: Unionida) and an evaluation of mitigation strategies. We found that drought studies were concentrated mostly in the USA, with a focus on the Unionidae family. Topics ranged from the physiological effects of high temperatures, emersion, and hypoxia/anoxia to behavioural and reproductive consequences of drought and the implications for biotic interactions and ecosystem services. Studies spanned all levels of biological organization, from individual responses to population- and community-level impacts and ecosystem-wide effects. We identified several knowledge gaps, including a paucity of trait-based evaluation of drought consequences, limited understanding of thermal and desiccation tolerance at the species level, and the synergistic effects of multiple drought stressors on mussels. Although we found many studies provided suggestions concerning management of populations, habitat conditions, and anthropogenic water use, a systematic approach and testing of recommended mitigation strategies is largely lacking, creating challenges for managers aiming to conserve freshwater mussel communities and populations in light of climate change.

Diurnal patterns of spatial stream temperature variations reveal the need for integrating thermal heterogeneity in riverscape habitat restoration.

Longer durations of warmer weather, altered precipitation, and modified streamflow patterns driven by climate change are expected to impair ecosystem resilience, exposing freshwater ecosystems and their biota to a severe threat worldwide. Understanding the spatio-temporal temperature variations and the processes governing thermal heterogeneity within the riverscape are essential to inform water management and climate adaptation strategies. We combined UAS-based imagery data of aquatic habitats with meteorological, hydraulic, river morphology and water quality data to investigate how key factors influence spatio-temporal stream heterogeneity on a diurnal basis within different thermal regions of a large recently restored Danube floodplain. Diurnal temperature ranges of aquatic habitats were larger than expected and ranged between 14.2 and 28.0 °C (mean = 20.7 °C), with peak median temperatures (26.1 °C) around 16:00 h. The observed temperature differences in timing and amplitude among thermal regions were unexpectedly high and created a mosaic pattern of temperature heterogeneity. For example, cooler groundwater-influenced thermal regions provided several cold water patches (CWP, below 19.0 °C) and potential cold water refuges (CWRs) around 12:00 h, at the time when other habitats were warmer than 21.0 °C, exceeding the ecological threshold (20.0 °C) for key aquatic species. Within the morphological complexity of the restored floodplain, we identified groundwater influence, shading and river morphology as the key processes driving thermal riverscape heterogeneity. Promoting stream thermal refuges will become increasingly relevant under climate change scenarios, and river restoration should consider both measures to physically prevent habitat from excessive warming and measures to improve connectivity that meet the temperature requirements of target species for conservation. This requires restoring mosaics of complex and dynamic temperature riverscapes.

Critical swimming speed of brown trout (Salmo trutta) infested with freshwater pearl mussel (Margaritifera margaritifera) glochidia and implications for artificial breeding of an endangered mussel species.

Unionid freshwater mussels need to attach to a host fish for completion of their life cycle. It remains unclear whether the relationship between these mussels and their host fishes can be considered parasitic, mutualistic, or commensal. Herein, we studied the effects of Margaritifera margaritifera infestation on Salmo trutta, the most important host of this endangered mussel species in Central Europe. Glochidial load of host fish increased with increasing glochidial concentration, but the highest ratios of encysted glochidia to exposed glochidia were found at low concentration (15,000 glochidia L(-1)) during infestation. Host fish mortality occurred at infestation rates of ~350 glochidia per g fish weight and was highest (60%) at the highest infestation rates (~900 glochidia per g fish weight). On a sublethal level, swimming performance of hosts was inversely related to infestation rates, with infestation of ~900 glochidia per g fish weight reducing critical swimming speed of S. trutta significantly by ~20% compared to infestation with 6 glochidia per g fish weight. The high mortality and the impaired swimming capability of highly infested hosts indicate a parasitic interaction between M. margaritifera and its host. For conservation and reintroduction of M. margaritifera via glochidia-infested S. trutta, we recommend glochidial loads of 5-100 glochidia per g fish weight, while for artificial breeding of juvenile M. margaritifera under laboratory conditions, higher infestation rates of up to 300 glochidia per g fish weight are ideal to balance high yields of mussels and welfare of host fishes.