ABSTRACT
The European eel (Anguilla anguilla) is a catadromous fish that spawns in the Sargasso Sea. As larvae, eels cross the Atlantic Ocean and reach the continental slope of Europe, where they ...metamorphose into post‐larval glass eels. These reach the continent, where some enter fresh water, some remain in marine waters, and others move between fresh and marine waters. After 5–25 years, as adult silver eels, they migrate back from fresh water to the Sargasso Sea to spawn and die. The glass eel stage is a critical step during which the eels cross the continental shelf and recruit to estuaries, where they facultatively transition to fresh water. Extensive research has been conducted to understand the behavioural mechanisms and environmental cues that aid and guide glass eels' migration. Glass eels follow odours and salinity gradients, they avoid light, and they change orientation and depth according to the tides. Recent work revealed that European glass eels also use Earth's magnetic field and lunar cues to orient. However, while we understand many aspects of their orientation behaviour, a unifying theory describing how glass eels migrate from the continental slope to fresh water is lacking. The goal of this review is to develop a comprehensive hypothesis on the migration of European glass eels, integrating previous knowledge on their orientation behaviour with recent findings on magnetic and celestial orientation. This review follows the journey of a hypothetical glass eel, describing the nature and the role of orientation cues involved at each step. I propose that, although glass eels have the sensory capacity to use multiple cues at any given time, their migration is based on a hierarchical succession of orientation mechanisms dictated by the physical properties of the environments that they occupy: (i) lunar and magnetic cues in pelagic water; (ii) chemical and magnetic cues in coastal areas; and (iii) odours, salinity, water current and magnetic cues in estuaries.
Anguillid eels grow in freshwater but spawn in the open ocean. The cues that guide eels over long distances to the spawning area are unknown. The Earth's magnetic field can provide directional and ...positional information and is likely used by catadromous eels during their spawning migration; as magnetosensitivity and compass orientation have been reported in eels. To test whether this is theoretically possible, we compared the migratory routes of five species of temperate eels that undertake long migrations with the geomagnetic field of their distribution/spawning areas. We found that, regardless of the species and although routes are different between life stages, larvae of those species always drift along paths of increasing magnetic inclination and intensity, while adults follow reverse gradients. This is consistent with an imprinting/retracing hypothesis. We propose a general navigation mechanism based on larvae imprinting on a target magnetic intensity (or inclination) at the hatching area and on the intensity (or inclination) gradient during larval drift. Years later, adults retrace the magnetic route by following the gradient of decreasing total intensity (or inclination) values that occurs towards lower latitudes. As they reach the target value, adults switch to compass orientation to stay on the target isoline and reach the spawning area. The proposed mechanism fits for all temperate eels examined. Knowledge about navigational strategies of eels is important to evaluate the effectiveness of management strategies that involve stocking of juveniles displaced from one area to another to rebuild local populations.
Sources of anthropogenic noise in the ocean have temporal and directional characteristics. Characterizing the soundscape requires identifying the directionality of the sources of noise in addition to ...the non-directional sound pressure. An underwater acoustic vector sensor (AVS) can be used to provide the directionality of incoming noise, and the concomitant sound pressure. We present an analysis of measurements from an AVS deployed in a Norwegian fjord in which there is frequent commercial ship traffic. We assessed the directionality of various known and unknown noise sources and used it to interpret the associated sound pressure. The fjord soundscape consists of time-varying noise directionality and intensity from anthropogenic sources, especially shipping activity. This case study highlights the benefits of using information from an AVS to assess noise directionality in a soundscape.
The number and size of offshore wind (OW) turbines is increasing rapidly. OW turbines produce continuous, low-frequency noise that could impact marine fish dispersing/migrating through the ...facilities. Any such impact would be relevant for larval stages, which have limited possibility to swim away from OW facilities. If directional movement of fish larvae at sea is impacted by low-frequency continuous sound is unknown. We observe the behavior of Atlantic cod larvae (N = 89) in response to low-frequency sound while they are drifting in a Norwegian fjord inside transparent drifting chambers. We transmit 100 Hz continuous sound in the fjord, in the intensity range of OW turbines' operational noise, and measure the sound pressure and 3-D particle motion. Half of the larvae (N = 45) are exposed to low-frequency (100 Hz) continuous sound, while the other half (N = 44) are observed under the same conditions but without the sound. Exposure does not affect the routine and maximum swimming speeds or the turning behavior of the larvae. Control larvae orient to the northwest. In contrast, exposed larvae orient towards the source of low-frequency sound and particle motion. This provides a basis to assess how OW might impact dispersal in this species.
The European eel (
) hatches in the Sargasso Sea and migrates to European and North African freshwater. As glass eels, they reach estuaries where they become pigmented. Glass eels use a tidal ...phase-dependent magnetic compass for orientation, but whether their magnetic direction is innate or imprinted during migration is unknown. We tested the hypothesis that glass eels imprint their tidal-dependent magnetic compass direction at the estuaries where they recruit. We collected 222 glass eels from estuaries flowing in different cardinal directions in Austevoll, Norway. We observed the orientation of the glass eels in a magnetic laboratory where the magnetic North was rotated. Glass eels oriented towards the magnetic direction of the prevailing tidal current occurring at their recruitment estuary. Glass eels use their magnetic compass to memorize the magnetic direction of tidal flows. This mechanism could help them to maintain their position in an estuary and to migrate upstream.
The European eel (
) has one of the longest migrations in the animal kingdom. It crosses the Atlantic Ocean twice during its life history, migrating between the spawning area in the Sargasso Sea and ...Europe, where it is widely distributed. The leptocephalus larvae drift with the Gulf Stream and other currents for more than a year and metamorphose into glass eels when they arrive on the continental shelf and move toward coastal areas. The mechanisms underlying glass eel orientation toward the coast and into freshwater systems are poorly known. However, anguillid eels, including the glass eel life stage, have a geomagnetic sense, suggesting the possibility that they use Earth's magnetic field to orient toward the coast. To test this hypothesis, we used a unique combination of laboratory tests and in situ behavioral observations conducted in a drifting circular arena. Most (98%) of the glass eels tested in the sea exhibited a preferred orientation that was related to the tidal cycle. Seventy-one percent of the same eels showed the same orientation during ebb tide when tested in the laboratory under a manipulated simulated magnetic field in the absence of any other cue. These results demonstrate that glass eels use a magnetic compass for orientation and suggest that this magnetic orientation system is linked to a circatidal rhythm.
Links between the lunar cycle and the life cycle (migration patterns, locomotor activity, pulses in recruitment) of the European eel (
) are well documented. In this study, we hypothesized that the ...orientation of glass eels at sea is related to the lunar cycle. The European eel hatches in the Sargasso Sea and migrates across the Atlantic Ocean towards Europe. Upon reaching the continental shelf, the larvae metamorphose into glass eels and migrate up the estuaries, where some individuals colonize freshwater habitats. How glass eels navigate pelagic waters is still an open question. We tested the orientation of 203 glass eels in a transparent circular arena that was drifting
during the daytime, in the coastal Norwegian North Sea, during different lunar phases. The glass eels swimming at sea oriented towards the azimuth of the moon at new moon, when the moon rose above the horizon and was invisible but not during the other moon phases. These results suggest that glass eels could use the moon position for orientation at sea and that the detection mechanism involved is not visual. We hypothesize a possible detection mechanism based on global-scale lunar disturbances in electrical fields and discuss the implications of lunar-related orientation for the recruitment of glass eels to estuaries. This behaviour could help glass eels to reach the European coasts during their marine migration.
Atlantic haddock (Melanogrammus aeglefinus) is a commercially important species of gadoid fish. In the North Sea, their main spawning areas are located close to the northern continental slope. Eggs ...and larvae drift with the current across the North Sea. However, fish larvae of many taxa can orient at sea using multiple external cues, including the Earth's magnetic field. In this work, we investigated whether haddock larvae passively drift or orient using the Earth's magnetic field. We observed the behavior of 59 and 102 haddock larvae swimming in a behavioral chamber deployed in the Norwegian North Sea and in a magnetic laboratory, respectively. In both in situ and laboratory settings, where the magnetic field direction was modified, haddock larvae significantly oriented toward the northwest. We conclude that haddock larvae orientation at sea is guided by a magnetic compass mechanism. These results have implications for retention and dispersal of pelagic haddock larvae.
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•Atlantic haddock larvae drift with the current and spread across the North Sea•In this area, larvae swimming in situ orient to the northwest•In a magnetic laboratory, larvae orient to the same direction, the magnetic northwest•Haddock larvae have a magnetic compass that they use to orient at sea
Piscine Behavior; Geomagnetism; Ichthyology
Abstract
Stakeholders need scientific advice on the environmental impacts of offshore wind (OW) before the facilities are installed. The utility of conventional environmental monitoring methods as a ...basis for forecasting OW impacts is limited because they do not explain the causes of the observed effects. We propose a multistep approach, based on process-oriented hypothesis testing, targeted monitoring and numerical modeling, to answer key stakeholder questions about planning an OW facility: Q1—Where do we place future OW farms so that impacts on the ecosystem are minimized? Q2—Which species and ecosystem processes will be impacted and to what degree? Q3—Can we mitigate impacts and, if so, how? and Q4—What are the risks of placing an OW facility in one location vs. another? Hypothesis testing can be used to assess impacts of OW facilities on target species-ecological process. This knowledge is transferable and is broadly applicable, a priori, to assess suitable locations for OW (Q1). Hypothesis testing can be combined with monitoring methods to guide targeted monitoring. The knowledge generated can identify the species/habitats at risk (Q2), help selecting/developing mitigation measures (Q3), and be used as input parameters for models to forecast OW impacts at a large spatial scale (Q1; Q4).
To orient themselves in their environment, animals integrate a wide array of external cues, which interact with several internal factors, such as personality. Here, we describe a behavioral protocol ...designed for the study of the influence of zebrafish personality on their orientation response to multiple external environmental cues, specifically water currents and magnetic fields. This protocol aims to understand whether proactive or reactive zebrafish display different rheotactic thresholds (i.e., the flow speed at which the fish start swimming upstream) when the surrounding magnetic field changes its direction. To identify zebrafish with the same personality, fish are introduced in the dark half of a tank connected with a narrow opening to a bright half. Only proactive fish explore the novel, bright environment. Reactive fish do not exit the dark half of the tank. A swimming tunnel with low flow rates is used to determine the rheotactic threshold. We describe two setups to control the magnetic field in the tunnel, in the range of the earth's magnetic field intensity: one that controls the magnetic field along the flow direction (one dimension) and one that allows a three-axial control of the magnetic field. Fish are filmed while experiencing a stepwise increase of the flow speed in the tunnel under different magnetic fields. Data on the orientation behavior are collected through a video-tracking procedure and applied to a logistic model to allow the determination of the rheotactic threshold. We report representative results collected from shoaling zebrafish. Specifically, these demonstrate that only reactive, prudent fish show variations of the rheotactic threshold when the magnetic field varies in its direction, while proactive fish do not respond to magnetic field changes. This methodology can be applied to the study of magnetic sensitivity and rheotactic behavior of many aquatic species, both displaying solitary or shoaling swimming strategies.
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