Alien species and climate change drive shifts in a riverine fish community and trait compositions over 35 years.

Alien fish substantially impact aquatic communities. However, their effects on trait composition remain poorly understood, especially at large spatiotemporal scales. Here, we used long-term biomonitoring data (1984-2018) from 31 fish communities of the Rhine river in Germany to investigate compositional and functional changes over time. Average total community richness increased by 49 %: it was stable until 2004, then declined until 2010, before increasing until 2018. Average abundance decreased by 9 %. Starting from 198 individuals/m2 in 1984 abundance largely declined to 23 individuals/m2 in 2010 (-88 %), and then consequently increased by 678 % up to 180 individuals/m2 until 2018. Increases in abundance and richness starting around 2010 were mainly driven by the establishment of alien species: while alien species represented 5 % of all species and 0.1 % of total individuals in 1993, it increased to 30 % (7 species) and 32 % of individuals in 2018. Concomitant to the increase in alien species, average native species richness and abundance declined by 26 % and 50 % respectively. We identified increases in temperature, precipitation, abundance and richness of alien fish driving compositional changes after 2010. To get more insights on the impacts of alien species on fish communities, we used 12 biological and 13 ecological traits to compute four trait metrics each. Ecological trait dispersion increased before 2010, probably due to diminishing ecologically similar native species. No changes in trait metrics were measured after 2010, albeit relative shares of expressed trait modalities significantly changing. The observed shift in trait modalities suggested the introduction of new species carrying similar and novel trait modalities. Our results revealed significant changes in taxonomic and trait compositions following alien fish introductions and climatic change. To conclude, our analyses show taxonomic and functional changes in the Rhine river over 35 years, likely indicative of future changes in ecosystem services.

The conservation paradox of critically endangered fish species: Trading alien sturgeons versus native sturgeon reintroduction in the Rhine-Meuse river delta.

Sturgeons rank among the most endangered vertebrates in the world. Yet, the dwindling of wild sturgeon populations stands in stark contrast to their thriving status in aquaculture. Moreover, through the exotic pet trade, sturgeons are introduced outside their natural ranges where they may compete and hybridize with native species and transmit parasites and diseases. Here, we present an in-depth inventory of alien sturgeons in the delta of the rivers Rhine and Meuse, because several countries consider reintroduction of the native, critically endangered European sturgeon (Acipenser sturio). Our study is based on (a) an inventory of the industry of sturgeon cultivation; (b) reports on spread of alien sturgeons; (c) an analysis of pathways for introduction and spread; and (d) a risk assessment using the Harmonia+ protocol. In total, 11 alien Acipenseriformes (sturgeons and paddlefishes) were traded across an intricate network of >1000 distribution points in the Netherlands, Belgium, and Germany. Circa 2500 alien sturgeons were reported from 53 angling ponds and 64 other lakes and ponds, whereas circa 500 alien sturgeons were reported widespread across hydrologically connected waters. Species that posed the highest risk of introduction, establishment and spread are Siberian sturgeon (A. baerii), Russian sturgeon (A. gueldenstaedtii) and Sterlet (A. ruthenus). We recommend to implement stringent trade regulations and practical solutions to prevent spread of alien sturgeons. Measures must preferably be taken at the spatial scale of river basins.

Sex differentiation in seasonal distribution of the starry smooth-hound Mustelus asterias.

This mark-recapture study of starry smooth-hound Mustelus asterias tagged during the summer months near the Dutch coast demonstrates a large-scale spatial sex differentiation in their circannual migration patterns and small-scale spatial sex differentiation during summer. Overwintering occurs in the North Sea, English Channel and Bay of Biscay, with significantly more males in the Northern North Sea and more females in the Bay of Biscay. During summer, sheltered sea arms off the Dutch coast were almost exclusively used by adult females. In subsequent summers post-release, both sexes were mostly confined to the Southern North Sea, suggesting philopatry.

Functional feeding traits as predictors of invasive success of alien freshwater fish species using a food-fish model.

Invasions of Ponto-Caspian fish species into north-western European river basins accelerated since the opening of the Rhine-Main-Danube Canal in 1992. Since 2002, at least five Ponto-Caspian alien fish species have arrived in The Netherlands. Four species belong to the Gobiidae family (Neogobius fluviatilis, Neogobius melanostomus, Ponticola kessleri, and Proterorhinus semilunaris) and one to the Cyprinidae family (Romanogobio belingi). These species are expected to be potentially deleterious for the populations of four native benthic fish species: Gobio gobio (Cyprinidae), Barbatula barbatula (Nemacheilidae), Cottus perifretum, and C. rhenanus (Cottidae). Invasion success may be dependent on competitive trophic interactions with native species, which are enabled and/or constrained by feeding-related morphological traits. Twenty-two functional feeding traits were measured in nine species (in total 90 specimens). These traits were quantitatively linked to the mechanical, chemical and behavioral properties of a range of aquatic resource categories, using a previously developed food-fish model (FFM). The FFM was used to predict the trophic profile (TP) of each fish: the combined capacities to feed on each of the resource types. The most extreme TPs belonged to three alien species, indicating that they were most specialized among the studied species. Of these three, only P. kessleri overlapped with the two native Cottus species, indicating potential trophic competition. N. fluviatilis and R. belingi did not show any overlap, indicating that there is low trophic competition. The two remaining alien goby species (N. melanostomus and P. semilunaris) had average TPs and could be considered generalist feeders. They overlapped with each other and with G. gobio and B. barbatula, indicating potential trophic competition. This study suggests that both generalist and specialist species can be successful invaders. Since the FFM predicts potential interactions between species, it provides a tool to support horizon scanning and rapid risk assessments of alien species.

Disentangling the complexity of tropical small-scale fisheries dynamics using supervised Self-Organizing Maps.

Tropical small-scale fisheries are typical for providing complex multivariate data, due to their diversity in fishing techniques and highly diverse species composition. In this paper we used for the first time a supervised Self-Organizing Map (xyf-SOM), to recognize and understand the internal heterogeneity of a tropical marine small-scale fishery, using as model the fishery fleet of San Pedro port, Tabasco, Mexico. We used multivariate data from commercial logbooks, including the following four factors: fish species (47), gear types (bottom longline, vertical line+shark longline and vertical line), season (cold, warm), and inter-annual variation (2007-2012). The size of the xyf-SOM, a fundamental characteristic to improve its predictive quality, was optimized for the minimum distance between objects and the maximum prediction rate. The xyf-SOM successfully classified individual fishing trips in relation to the four factors included in the model. Prediction percentages were high (80-100%) for bottom longline and vertical line + shark longline, but lower prediction values were obtained for vertical line (51-74%) fishery. A confusion matrix indicated that classification errors occurred within the same fishing gear. Prediction rates were validated by generating confidence interval using bootstrap. The xyf-SOM showed that not all the fishing trips were targeting the most abundant species and the catch rates were not symmetrically distributed around the mean. Also, the species composition is not homogeneous among fishing trips. Despite the complexity of the data, the xyf-SOM proved to be an excellent tool to identify trends in complex scenarios, emphasizing the diverse and complex patterns that characterize tropical small scale-fishery fleets.

Evidence for ontogenetically and morphologically distinct alternative reproductive tactics in the invasive Round Goby Neogobius melanostomus.

Alternative reproductive tactics are characterized by the occurrence of discrete alternative morphs that differ in behavioural, morphological and physiological traits within the same sex. Although much effort has been made to describe the behaviour, morphology and physiology of such alternative morphs, less effort has been invested investigating how much overlap there is in the characteristics of such morphs in natural populations. We studied random population samples of the invasive Round Goby Neogobius melanostomus from five different localities in the river Rhine system in the Netherlands. We found two morphologically and physiologically distinct male morphs which likely represent alternative reproductive tactics. Almost all mature males under 9.35 cm total length had a gonadosomatic index > 3%, suggestive of a sneaker tactic, while nearly all males above 9.35 cm has a gonadosomatic index of < 3%, suggestive of a parental tactic. Cheek size and eye diameter alone were sufficient to distinguish the two morphs. Gonads had a different relationship with size in the two morphs, indicating separate growth trajectories. The gonad mass of sneaker morphs would be ca. 7.5 times as high as the gonad mass of parental morphs of the same total length after extrapolation. Few (9%) intermediates were found, suggesting that the expression of alternative reproductive tactics is determined before the first breeding season. This contrasts with studies on other goby species, which show evidence of plastic tactics that can be affected by social circumstances. We conclude that it is possible to distinguish two alternative male morphs in the Dutch Round Goby population using morphological measurements alone. Although behavioural observations are needed to provide conclusive evidence, the difference in GSI between these morphs indicates that these morphs reflect alternative reproductive tactics.

Serial correlation and inter-annual variability in relation to the statistical power of monitoring schemes to detect trends in fish populations.

We studied the effects of inter-annual variability and serial correlation on the statistical power of monitoring schemes to detect trends in biomass of bream (Abramis brama) in Lake Veluwemeer (The Netherlands). In order to distinguish between 'true' system variability and sampling variability we simulated the development of the bream population, using estimates for population structure and growth, and compared the resulting inter-annual variabilities and serial correlations with those from field data. In all cases the inter-annual variability in the field data was larger than in simulated data (e.g. for total biomass of all assessed bream sigma = 0.45 in field data, and sigma = 0.03-0.14 in simulated data) indicating that sampling variability decreased statistical power for detecting trends. Moreover, sampling variability obscured the inter-annual dependency (and thus the serial correlation) of biomass, which was expected because in this long-lived population biomass changes are buffered by the many year classes present. We did find the expected serial correlation in our simulation results and concluded that good survey data of long-lived fish populations should show low sampling variability and considerable inter-annual serial correlation. Since serial correlation decreases the power for detecting trends, this means that even when sampling variability would be greatly reduced, the number of sampling years to detect a change of 15%.year(-1) in bream populations (corresponding to a halving or doubling in a six-year period) would in most cases be more than six. This would imply that the six-year reporting periods that are required by the Water Framework Directive of the European Union are too short for the existing fish monitoring schemes.