Individual based models for the simulation of fish movement near barriers: Current work and future directions.

River fragmentation is an increasing issue for water managers and conservationists. Barriers such as dams interfere with freshwater fish migration, leading to drastic population declines. While there are a range of widely implemented mitigation approaches, e.g. fish passes, such measures are often inefficient due to suboptimal operation and design. There is increasing need to be able to assess mitigation options prior to implementation. Individual based models (IBMs) are a promising option. IBMs can simulate the fine-scale movement of individual fish within a population as they attempt to find a fish pass, incorporating movement processes themselves. Moreover, IBMs have high transferability to other sites or conditions (e.g. changing mitigation, change in flow conditions), making them potentially valuable for freshwater fish conservation yet their application to the fine-scale movement of fish past barriers is still novel. Here, we present an overview of existing IBMs for fine-scale freshwater fish movement, with emphasis on study species and the parameters driving movement in the models. In this review, we focus on IBMs suitable for the simulation of fish tracks as they approach or pass a single barrier. The selected IBMs for modelling fine-scale freshwater fish movement largely focus on salmonids and cyprinid species. IBMs have many applications in the context of fish passage, such as testing different mitigation options or understanding processes behind movement. Existing IBMs include movement processes such as attraction and rejection behaviours, as reported in literature. Yet some factors affecting fish movement e.g. biotic interactions are not covered by existing IBMs. As the technology available for fine scale data collection continues to advance, such as increasing data linking fish behaviour to hydraulics, IBMs could become a more common tool in the design and implementation of fish bypass structures.

Glass eel (Anguilla anguilla L. 1758) feeding behaviour during upstream migration in an artificial waterway.

The transition from marine to fresh water is a challenging task for juvenile eels. This critical step in the early eels' life is preceded by a metamorphosis from the oceanic larval to the continental glass eel stage, requiring major energy-demanding morphological, physiological and behavioural modifications during which time these animals do not feed. The success of the glass eels' inland migration after metamorphosis will largely depend on remaining energy levels, which can be supplemented only by resuming food uptake. Although it is crucial for their survival and the maintenance of the population, the feeding behaviour of glass eels is still an understudied aspect of the eels' complex life cycle. Many uncertainties about the phenology, diet, potential prey preferences and their relation with migration modus (migratory vs. sedentary) still remain. In this study, the authors analysed the stomach and gut contents of 458 European glass eels (Anguilla anguilla L. 1758) captured in a drainage canal connecting a small mesotidal estuary with an adjacent polder area during the spring migration seasons of 2016 and 2017. They demonstrated that although glass eels started feeding briefly upon arrival in the estuary, food uptake for early arrivals was restricted to a minority that sparsely feed on detritus and some worm-like benthic invertebrates. Along the season, food uptake intensified eventually engaging all glass eels and their dietary palette diversified including a wide array of planktonic and benthic organisms. Crustacean plankton (mainly cyclopoid copepods) was an important part of the glass eel diet, whereas benthic oligochaetes were less abundant as food source in spite of their high presence in the sediments. No clear differences in feeding behaviour could be observed between migratory and sedentary glass eels. This study showed that glass eels can use highly artificial and dynamic drainage canals as feeding ground during their critical marine/freshwater transition. This outcome is also a plea to improve the accessibility of alternative (unnatural) migration routes between the ocean and suitable freshwater growth habitats for the European eel.

Finding navigation cues near fishways.

Many fish species depend on migration for various parts of their life cycle. Well-known examples include diadromous fish such as salmon and eels that need both fresh water and salt water to complete their life cycle. Migration also occurs within species that depend only on fresh water. In recent decades, anthropogenic pressures on freshwater systems have increased greatly, and have resulted, among other effects, in drastic habitat fragmentation. Fishways have been developed to mitigate the resulting habitat fragmentation, but these are not always effective. To improve fishway efficiency, the variety of navigation cues used by fish must be better understood: fish use a multitude of sensory inputs ranging from flow variables to olfactory cues. The reaction of a fish is highly dependent on the intensity of the cue, the fish species involved, and individual traits. Recently developed monitoring technologies allow us to gain insights into different combinations of environmental and physiological conditions. By combining fish behavioural models with environmental models, interactions among these components can be investigated. Several methods can be used to analyse fish migration, with state-space models, hidden Markov models, and individual-based models potentially being the most relevant since they can use individual data and can tie them to explicit spatial locations within the considered system. The aim of this review is to analyse the navigational cues used by fish and the models that can be applied to gather knowledge on these processes. Such knowledge could greatly improve the design and operation of fishways for a wider range of fish species and conditions.

Individual variation in the habitat selection of upstream migrating fish near a barrier.

BACKGROUND: Migration is a vital element of the life cycle of many freshwater fish species but is increasingly hampered globally by riverine barriers. Fish passes are a common approach to enable migration past barriers but are often ineffective. More knowledge is required on fish behaviour as they approach barriers such as habitat preferences. METHODS: We evaluate the habitat selection of two upstream migrating fish species, barbel Barbus barbus and grayling Thymallus thymallus, at a hydropower plant in southern Germany, considering individual variation and population trends. Fish were tracked via fine-scale 2D acoustic telemetry in 2018 during their spawning migration. Step selection functions were used to evaluate selection of hydraulic parameters by the fish for a time step of 20 s. Exploratory models were built via model selection for each individual fish, to evaluate the extent of individual variation in model structure. A population model was developed for each species by averaging coefficients from individual models to describe general trends. The extent of individual variation was determined and confidence intervals for the population model coefficients were calculated. RESULTS: Fish varied greatly in individual model structure though common terms were apparent in both species, such as depth, flow velocity, the angular difference between fish and velocity, and the logarithm of the step length. Final population models for barbel included several parameters describing habitat selection and displacement. Barbel selected for faster flows, deeper water, and higher spatial velocity gradients. In addition, they selected to move more with the flow than against. Interactions were also present between habitat parameters, suggesting selection is context dependent. Barbel movement speed also changed with depth, flow velocity and spatial velocity gradient. With grayling, terms often had contrasting effects among individuals and thus general trends could not be distinguished for most terms. CONCLUSION: Our findings demonstrate habitat selection by upstream migrating fish approaching a fish pass and differences in individual selection which may have an impact on barrier management. Step selection functions are a promising approach and can provide useful insight into habitat selection and movement by migrating freshwater fish in an altered river system.

Tidal and circadian patterns of European eel during their spawning migration in the North Sea and the English Channel.

Technological advances in tracking methods enable the mapping of anguillid eel migration routes from continental habitats to their spawning sites in the ocean. However, the behaviour and orientation abilities of anguillids are still poorly understood, and have only rarely been studied on the continental shelf. Here we present the results of a study into the vertical and horizontal movement behaviour of 42 European eels (Anguilla anguilla L.) tagged with electronic tags that migrated through the North Sea and English Channel towards and into the Atlantic Ocean during their spawning migration. We used actograms, periodograms and linear mixed effects models to determine the periodicity and significance of the timing and pattern of vertical movement and activity. Overall, eels had a complex behavioural repertoire that included classical diel vertical migration (DVM), reverse DVM and vertical movement behaviours that synchronized with tidal patterns. All of the eels that were tracked showed one or more of these behaviours during their time at liberty, and many exhibited all of them. We also observed that the eels had a higher horizontal migration speed when the current in the favourable direction was stronger. This, together with the vertical movement synchronized with the tides, suggests the eels adopt selective tidal stream transport. Finally, tracked eels had a higher vertical movement range at night compared to daytime. We hypothesize that these behaviours are driven by bio-energetic efficient movement, navigation and predator avoidance.

Mapping silver eel migration routes in the North Sea.

Recent developments in tracking technology resulted in the mapping of various marine spawning migration routes of the European eel (Anguilla anguilla). However, migration routes in the North Sea have rarely been studied, despite many large European rivers and hence potential eel growing habitat discharge into the North Sea. In this study, we present the most comprehensive map to date with migration routes by silver European eels in the North Sea and document for the first time successful eel migration through the English Channel. Migration tracks were reconstructed for 42 eels tagged in Belgium and 12 in Germany. Additionally, some eels moved up north to exit the North Sea over the British Isles, confirming the existence of two different routes, even for eels exiting from a single river catchment. Furthermore, we observed a wide range in migration speeds (6.8-45.2 km day-1). We hypothesize that these are likely attributed to water currents, with eels migrating through the English Channel being significantly faster than eels migrating northward.

Unimodal head-width distribution of the European eel (Anguilla anguilla L.) from the Zeeschelde does not support disruptive selection.

Since the early 20th century, European eels (Anguilla anguilla L.) have been dichotomously classified into 'narrow' and 'broad' heads. These morphs are mainly considered the result of a differential food choice, with narrow heads feeding primarily on small/soft prey and broad heads on large/hard prey. Yet, such a classification implies that head-width variation follows a bimodal distribution, leading to the assumption of disruptive selection. We investigated the head morphology of 272 eels, caught over three consecutive years (2015-2017) at a single location in the Zeeschelde (Belgium). Based on our results, BIC favored a unimodal distribution, while AIC provided equal support for a unimodal and a bimodal distribution. Notably, visualization of the distributions revealed a strong overlap between the two normal distributions under the bimodal model, likely explaining the ambiguity under AIC. Consequently, it is more likely that head-width variation followed a unimodal distribution, indicating there are no disruptive selection pressures for bimodality in the Zeeschelde. As such, eels could not be divided in two distinct head-width groups. Instead, their head widths showed a continuum of narrow to broad with a normal distribution. This pattern was consistent across all maturation stages studied here.