The biological invasion of an apex predator (Silurus glanis) amplifies PCB transfer in a large lake food web.

Invasive species can affect food web structure possibly modifying the transfer of pollutants in ecosystems but this facet of biological invasion remains largely unexplored. We examined how trophic and ontogenetic characteristics of the invasive European catfish could differ from its native counterpart, the Northern pike, possibly resulting in the amplification of PCB transfer to the higher trophic levels in a large lake food web. The PCB contents of catfish and pike were on average low (Æ©7 PCBi 42.4 ± 38.6 ng g-1 ww and 37.9 ± 49.4 ng g-1 ww respectively) and dominated by PCB153 (~35 % of the PCB contamination). Only the largest pike (126 cm) slightly exceeded the European sanitary threshold of 125 ng g-1 ww Æ©6 PCBi-NDL. Both species increased in trophic position with body size while catfish had clearly higher littoral reliance than pike indicating they exploited complementary trophic niches. PCB biomagnification was identified only for catfish (PCB153, Æ©7 PCBi) leading to trophic magnification factor of ~5. PCB ontogenetic bioaccumulation was pervasive for catfish (PCB101, PCB118, PCB153, PCB138 and Æ©7 PCBi) and identified for pike only regarding PCB101. The derived size accumulation factors (~1.02) indicated a size-doubling PCB contamination of ~40 cm for catfish. This finding suggested that catfish would exceed the European sanitary threshold at body size larger than 168 cm possibly constraining their commercial exploitation. Our results highlighted that the invasive catfish was a littoral-oriented apex predator occupying an alternative trophic niche as compared to pike thereby modifying the lake food web structure that resulted in an enhancement of PCB transfer to higher trophic levels. The biomagnification and ontogenetic bioaccumulation of catfish underlined the impact of this biological invasion on the fate of PCB in the ecosystem. Finally, the remarkable inter-individual PCB contamination suggested variable inter-individual PCB exposure likely associated to localized hotspots of PCB contamination in the lake.

Overwintering aggregation patterns of European catfish Silurus glanis.

Animal aggregation, particularly in large-bodied species, is both a fascinating and intriguing phenomenon. Here we analyzed the overwintering behavior of the European catfish, Silurus glanis Linnaeus, 1758, the largest freshwater fish in Europe. By tracking 47 subadults and adults in a shallow lake in southeastern France, we reported a consistent aggregative behavior across four successive winters. By implementing time series analysis and Cox proportional hazard models, we investigated the dynamics of these aggregations (formation, stability, dislocation), and the factors that govern it, whether external (temperature, time of the day) or specific to the fish (size, key individuals). These aggregations lasted 1.5-2 months and mainly took place in a single small 4 m-deep area whose environmental conditions (temperature, oxygen, substrate) did not differ from other parts of the lake. In some periods during winter, all tagged fish were aggregated, which suggests that a large proportion of the lake population gathered there. Low temperatures (below 9 °C) triggered the formation of aggregations. They became more stable with decreasing temperatures, while individuals more frequently left the aggregation, preferentially at dusk and at night, when temperatures increased. The largest individuals swam more frequently back and forth to the aggregation. Irrespective of their size, some individuals consistently arrived earlier in the aggregation in winter and left later. This predictable seasonal grouping of individuals and, more generally, the knowledge provided by such studies on how species use space have important operational value and are useful for species conservation as well as for species control.

Intraspecific diversity loss in a predator species alters prey community structure and ecosystem functions.

Loss in intraspecific diversity can alter ecosystem functions, but the underlying mechanisms are still elusive, and intraspecific biodiversity-ecosystem function (iBEF) relationships have been restrained to primary producers. Here, we manipulated genetic and functional richness of a fish consumer (Phoxinus phoxinus) to test whether iBEF relationships exist in consumer species and whether they are more likely sustained by genetic or functional richness. We found that both genotypic and functional richness affected ecosystem functioning, either independently or interactively. Loss in genotypic richness reduced benthic invertebrate diversity consistently across functional richness treatments, whereas it reduced zooplankton diversity only when functional richness was high. Finally, losses in genotypic and functional richness altered functions (decomposition) through trophic cascades. We concluded that iBEF relationships lead to substantial top-down effects on entire food chains. The loss of genotypic richness impacted ecological properties as much as the loss of functional richness, probably because it sustains "cryptic" functional diversity.

High predation of native sea lamprey during spawning migration.

Sea lamprey (Petromyzon marinus) is a unique jawless vertebrate among the most primitive of all living vertebrates. This migratory fish is endangered in much of its native area due to dams, overfishing, pollution, and habitat loss. An introduced predator, the European catfish (Silurus glanis), is now widespread in Western and Southern European freshwaters, adding a new threat for sea lamprey migrating into freshwater to spawn. Here, we use a new prototype predation tag coupled with RFID telemetry on 49 individuals from one of the largest sea lamprey European populations (Southwestern France) to quantify the risk of predation for adult sea lampreys during its spawning migration in rivers with large populations of European catfish. We found that at least 80% of tagged sea lampreys (39 among 49) were preyed upon within one month, and that 50% of the released lampreys were rapidly consumed on average 8 days after tagging. This very high predation rate suggests that the European catfish represents a supplementary serious threat of extirpation for the native sea lamprey population we studied. This threat is likely to happen throughout most of the native lamprey distribution area, as the European catfish is becoming established almost everywhere the sea lamprey is.

Within-individual trophic variability drives short-term intraspecific trait variation in natural populations.

Intraspecific trait variability (ITV) maintains functional diversity in populations and communities, and plays a crucial role in ecological and evolutionary processes such as trophic cascades or speciation. Furthermore, functional variation within a species and its populations can help buffer against harmful environmental changes. Trait variability within species can be observed from differences among populations, and between- and within individuals. In animals, ITV can be driven by ontogeny, the environment in which populations live and by within-individual specialization or variation unrelated to growth. However, we still know little about the relative strength of these drivers in determining ITV variation in natural populations. Here, we aimed to (a) measure the relative strength of between- and within-individual effects of body size on ITV over time, and (b) disentangle the trophic changes due to ontogeny from other sources of variability, such as the environment experienced by populations and individual preferences at varying temporal and spatial scales. We used as a model system the endangered marble trout Salmo marmoratus, a freshwater fish living in a restricted geographical area (<900 km2 ) that shows marked changes in diet through ontogeny. We investigated two trophic traits, trophic position and resource use, with stable isotopes (δ15 N and δ13 C), and followed over time 238 individually tagged marble trout from six populations to estimate the trophic changes between and within individuals through ontogeny at three different time-scales (short term: 3 months, medium term: 1 year and long term: 2 years). We found that the relative strength of between- and within-individual effects of body size on trophic position and resource use change strongly over time. Both effects played a similar role in ITV over medium- and long-term time-scales, but within-individual effects were significantly driving trophic variability over short-term scales. Apart from ontogenetic shifts, individuals showed variability in trophic traits as big as the variability estimated between populations. Overall, our results show how the relative strengths of ITV drivers change over time. This study evidences the crucial importance of considering effects of time-scales on functional variability at individual, population and species levels.

Environmental conditions and neutral processes shape the skin microbiome of European catfish (Silurus glanis) populations of Southwestern France.

Teleost fishes interact with diverse microbial communities, playing crucial functions for host fitness. While gut microbiome has been extensively studied, skin microbiome has been overlooked. Specifically, there is no assessment of the relative impact of host and environmental factors on microbiome variability as well as neutral processes shaping fish skin microbiome. Here, we assessed the skin microbiome of a Siluriforme, the European catfish (Silurus glanis) sampled in four sites located in Southwestern France. We assessed the relative roles of individual features (body size and genetic background), local environment and neutral processes in shaping skin microbiome. Catfish skin microbiome composition was distinct to that of other freshwater fish species previously studied with high abundances of Gammaproteobacteria and Bacteroidetes. We found no effect of catfish individual genotype and body size on the structure of its associated skin microbiome. Geographical location was the best catfish skin microbiome structure predictor, together with neutral models of microbiome assembly.

Variability of functional traits and their syndromes in a freshwater fish species (Phoxinus phoxinus): The role of adaptive and nonadaptive processes.

Functional traits can covary to form "functional syndromes." Describing and understanding functional syndromes is an important prerequisite for predicting the effects of organisms on ecosystem functioning. At the intraspecific level, functional syndromes have recently been described, but very little is known about their variability among populations and-if they vary-what the ecological and evolutionary drivers of this variation are. Here, we quantified and compared the variability in four functional traits (body mass, metabolic rate, excretion rate, and boldness), their covariations and the subsequent syndromes among thirteen populations of a common freshwater fish (the European minnow, Phoxinus phoxinus). We then tested whether functional traits and their covariations, as well as the subsequent syndromes, were underpinned by the phylogenetic relatedness among populations (historical effects) or the local environment (i.e., temperature and predation pressure), and whether adaptive (selection or plasticity) or nonadaptive (genetic drift) processes sustained among-population variability. We found substantial among-population variability in functional traits and trait covariations, and in the emerging syndromes. We further found that adaptive mechanisms (plasticity and/or selection) related to water temperature and predation pressure modulated the covariation between body mass and metabolic rate. Other trait covariations were more likely driven by genetic drift, suggesting that nonadaptive processes can also lead to substantial differences in trait covariations among populations. Overall, we concluded that functional syndromes are population-specific, and that both adaptive and nonadaptive processes are shaping functional traits. Given the pivotal role of functional traits, differences in functional syndromes within species provide interesting perspectives regarding the role of intraspecific diversity for ecosystem functioning.

The community and ecosystem consequences of intraspecific diversity: a meta-analysis.

Understanding the relationships between biodiversity and ecosystem functioning has major implications. Biodiversity-ecosystem functioning relationships are generally investigated at the interspecific level, although intraspecific diversity (i.e. within-species diversity) is increasingly perceived as an important ecological facet of biodiversity. Here, we provide a quantitative and integrative synthesis testing, across diverse plant and animal species, whether intraspecific diversity is a major driver of community dynamics and ecosystem functioning. We specifically tested (i) whether the number of genotypes/phenotypes (i.e. intraspecific richness) or the specific identity of genotypes/phenotypes (i.e. intraspecific variation) in populations modulate the structure of communities and the functioning of ecosystems, (ii) whether the ecological effects of intraspecific richness and variation are strong in magnitude, and (iii) whether these effects vary among taxonomic groups and ecological responses. We found a non-linear relationship between intraspecific richness and community and ecosystem dynamics that follows a saturating curve shape, as observed for biodiversity-function relationships measured at the interspecific level. Importantly, intraspecific richness modulated ecological dynamics with a magnitude that was equal to that previously reported for interspecific richness. Our results further confirm, based on a database containing more than 50 species, that intraspecific variation also has substantial effects on ecological dynamics. We demonstrated that the effects of intraspecific variation are twice as high as expected by chance, and that they might have been underestimated previously. Finally, we found that the ecological effects of intraspecific variation are not homogeneous and are actually stronger when intraspecific variation is manipulated in primary producers than in consumer species, and when they are measured at the ecosystem rather than at the community level. Overall, we demonstrated that the two facets of intraspecific diversity (richness and variation) can both strongly affect community and ecosystem dynamics, which reveals the pivotal role of within-species biodiversity for understanding ecological dynamics.

Catch-related and genetic outcome of adult northern pike Esox lucius stocking in a large river system.

Genetic introgression from stocked adult northern pike Esox lucius to a wild self-recruiting population was detected in a large river system and some stocked E. lucius survived up to two spawning seasons and dispersed over several kilometres in the river. Moreover, the catch rate of stocked E. lucius by anglers was low (9.6%), hence suggesting that the efficiency of stocking activity is questionable.

Adult Atlantic salmon have a new freshwater predator.

The Atlantic salmon (Salmo salar) is one of the world's most emblematic freshwater fish. Despite conservation and rehabilitation plans, populations of this species are dramatically declining due to human impacts such as habitat fragmentation, overfishing and water pollution. Owing to their large body size, anadromous adults were historically invulnerable to fish predation during their spawning period migration. This invulnerability has disappeared in Western Europe with the introduction of a new freshwater predator, the European catfish (Silurus glanis). Here we report how adults of Atlantic salmon are predated in the fishway of a large river of SW France, where the delayed and narrow passage created by the structure increases the probability of predator-prey encounter. We assessed predation risk by monitoring salmon and catfish in one fishway of the River Garonne, using video fish-counting from 1993 to 2016. We analysed the predation strategy of catfish using observations made with acoustic camera and RFID telemetry in 2016. Our results demonstrate a high predation rate (35%-14/39 ind.) on salmon inside the fishway during the 2016 spawning period migration. Our results suggest that a few specialized catfish individuals adapted their hunting behaviour to such prey, including their presence synchronized with that of salmon (i.e, more occurrences by the end of the day). Such results suggest that the spread of European catfish will potentially impact migration of anadromous species through anthropized systems.

"Freshwater killer whales": beaching behavior of an alien fish to hunt land birds.

The behavioral strategies developed by predators to capture and kill their prey are fascinating, notably for predators that forage for prey at, or beyond, the boundaries of their ecosystem. We report here the occurrence of a beaching behavior used by an alien and large-bodied freshwater predatory fish (Silurus glanis) to capture birds on land (i.e. pigeons, Columbia livia). Among a total of 45 beaching behaviors observed and filmed, 28% were successful in bird capture. Stable isotope analyses (δ(13)C and δ(15)N) of predators and their putative prey revealed a highly variable dietary contribution of land birds among individuals. Since this extreme behavior has not been reported in the native range of the species, our results suggest that some individuals in introduced predator populations may adapt their behavior to forage on novel prey in new environments, leading to behavioral and trophic specialization to actively cross the water-land interface.

Colossal aggregations of giant alien freshwater fish as a potential biogeochemical hotspot.

The ubiquity and fascinating nature of animal aggregations are widely recognised. We report here consistent and previously undocumented occurences of aggregations of a giant alien freshwater fish, the Wels catfish (Silurus glanis). Aggregative groups were on average composed of 25 (± 10 SD, ranging from 15 to 44) adults with estimated average total biomass of 651 kg (386 - 1132) and biomass density of 23 kg m(-2) (14 - 40). Aggregations always occurred within the same location. No foraging, reproductive or anti-predator behaviour were observed during the aggregations. A mass-balance model estimated that these colossal aggregations of an alien species can locally release, through excretion only, up to 70 mg P m(-2) h(-1) and 400 mg N m(-2) h(-1), potentially representing the highest biogeochemical hotspots reported in freshwater ecosystems and another unexpected ecological effect of alien species.

Trophic ecology of the pikeperch (Sander lucioperca) in its introduced areas: a stable isotope approach in southwestern France.

During the last decades, non-native predatory fish species have been largely introduced in European lakes and rivers, calling for detailed information on the trophic ecology of co-existing native and non-native predators. The present study describes the trophic ecology of the introduced pikeperch (Sander lucioperca) in two southwestern French rivers, using stable isotope analysis. Pikeperch could be categorized as a top-predator, and had a significantly higher trophic position (TP, mean+/-SE=4.2+/-0.1) compared to other predatory fish such as the native pike (Esox lucius, TP=3.7+/-0.1) and the introduced European catfish (Silurus glanis, TP=3.8+/-0.1). Most studies of resource use in freshwaters consider predatory fish as ecologically equivalent; however, this study showed that the pikeperch occupied a higher trophic niche compared to other predatory species in the Lot and Tarn rivers (Garonne River basin). This apparent specialization may thus have consequences upon interspecific relationships within the predatory guild and upon the functional organization of biological communities.

Contribution of anadromous fish to the diet of European catfish in a large river system.

Many anadromous fish species, when migrating from the sea to spawn in fresh waters, can potentially be a valuable prey for larger predatory fish, thereby efficiently linking these two ecosystems. Here, we assess the contribution of anadromous fish to the diet of European catfish (Silurus glanis) in a large river system (Garonne, southwestern France) using stable isotope analysis and allis shad (Alosa alosa) as an example of anadromous fish. Allis shad caught in the Garonne had a very distinct marine delta(13)C value, over 8 per thousand higher after lipid extraction compared to the mean delta(13)C value of all other potential freshwater prey fish. The delta(13)C values of European catfish varied considerably between these two extremes and some individuals were clearly specializing on freshwater prey, whereas others specialized on anadromous fish. The mean contribution of anadromous fish to the entire European catfish population was estimated to be between 53% and 65%, depending on the fractionation factor used for delta(13)C.

Co-occurrence patterns of some small-bodied freshwater fishes in southwestern France: implications for fish conservation and environmental management.

We assessed the influence of environmental variables (elevation, stream order, distance from source, catchment area, slope, stream width, and fish species richness) on the co-occurrence patterns of the minnow, the stone loach, and the gudgeon at the stream system scale. A total of 474 sites were classified according to the seven variables using the Self-Organizing Map (neural network), and three clusters were detected (k-means algorithm). The frequency of the various fish co-occurrence patterns was calculated for each cluster, and general linear modeling was used to specify the conditions that predict the occurrence of each species. Piedmont streams were more likely to support coexisting gudgeon and minnow populations because of higher probabilities of occurrence for both species. The higher co-occurrence frequency for the three species together in headwater streams resulted from lower occurrence frequencies in gudgeon and minnow. Focusing on areas that favor the co-occurrence of species may enhance the effectiveness of conservation projects.