Individual patterns of rhythmic swimming activity in Anguilla anguilla glass eels synchronised to water current reversal

The existence of distinct patterns of activity and swimming behaviour were tested in individual European glass eels by means of Bayesian inference mixture modelling. 36 glass eels were tagged using Visible Implant Elastomer and added to 36 untagged glass eels in February and April. Each group was presented with a change in water current direction every 6,2 h and videotaped during 2 weeks. Two hypotheses were tested: (i) all individuals display a similar pattern of behaviour within a tidal period, glass eels showing both positive and negative rheotaxis in opposite phase (M1 model) and (ii) individuals are distributed in two different groups, some glass eels swimming with and the others against the current (M2 model). Results showed that most glass eels displayed a positive or a negative rheotaxis and only a small number exhibited both behaviours. All swimming behaviours were synchronised to a change in current direction with a period close to the tidal one. Results are discussed in relation to synchronisation and migration behaviour.     

Annual variability in upstream migration of glass eels in a southern USA coastal watershed

We investigated the environmental factors that affected temporal variability of eel recruitment and upstream migration in a freshwater coastal river along the southeastern US. Glass eels Anguilla rostrata were collected through ichthyoplankton sampling in the lower Roanoke River, North Carolina. Monthly samples were taken from fixed stations from May 2001 through June 2003. There was no evidence of consistent seasonal migration patterns for glass eels in Roanoke River. From May through December in 2001, glass eels were captured only during August. In 2002, glass eels arrived in February and remained in ichthyoplankton samples through October, with the exception of samples from September. Peak catch occurred in March at 4.02 ± 1.2 and declined through June to 0.18 ± 0.07 (#/1,000 m³). By August, the mean density increased to 0.96 ± 0.82 and to 3.59 ± 2.77 by October. In 2003 from January through June, glass eels were captured only during February and March. Glass eels were routinely collected when river discharge rates were <150 m³ s⁻¹. River discharge rates >650 m⁻³ s⁻¹ resulted in no glass eels in our samples. Upstream migration during 2002 was not correlated with water temperature or related to lunar phase. Glass eel freshwater upstream migration was initiated when water temperatures exceeded a threshold range of 10°C to 15°C; however, glass eels continued to migrate when water temperatures approached 30°C. The overall negative effect of river discharge suggests that changes in the water release schedules of upstream hydroelectric facilities during glass eel migration could strongly influence their recruitment success.     

Dispersal of yellow phase Japanese eels <Emphasis Type="Italic">Anguilla japonica</Emphasis> after recruitment in the Kojima Bay-Asahi River system,Japan

The density, size and age distribution were investigated for 233 eels, Anguilla japonica, sampled in fresh and brackish water areas of the Kojima Bay-Asahi River system, Okayama, Japan, to evaluate the possible patterns of dispersal of eels that recruit to this area. Migratory histories of 183 eels were categorized into 5 types depending on the Sr and Ca concentrations in their otoliths: (1) brackish water residents (74 fish, 40.4%), which settled in saline water and remained until capture; (2) freshwater residents (46 fish, 25.1%), which settled in freshwater and remained until capture; (3) upstream shifters (3 fish, 1.6%), which settled in saline water and moved upstream into freshwater; (4) downstream shifters (53 fish, 29.0%), which settled in freshwater and moved downstream into saline water; (5) multiple habitat shifters (7 fish, 3.8%), which shifted their habitats between freshwater and saline water more than twice. For eels captured in the brackish water area, fish density decreased with distance in the downstream direction, while the size and age of eels increased. For eels captured in the freshwater area, size and age were greater than those in the upper-most brackish site. These observations suggest that eels in this system initially accumulate in the lower reaches of the river and then disperse in both upstream and downstream directions following their growth.     

Estuarine habitat preferences of <Emphasis Type="Italic">Anguilla australis</Emphasis> and <Emphasis Type="Italic">A. reinhardtii</Emphasis> glass eels as inferred from laboratory experiments

We tested the habitat preferences of Anguilla australis (shortfin) and A. reinhardtii (longfin) glass eels using circular tanks in an aquarium, containing four types of estuarine habitat (sand, mud, rocks/cobbles and seagrass). Shortfin eels either showed a tendency to occur in heterogeneous habitats, or in rocks/cobbles. Longfin glass eels showed a significant preference for rocks/cobbles in both experiments. Tests on shortfin and longfin glass eels in tanks with only rocks/cobbles available showed that eels were not clumped, indicating that individuals select habitat for re-settlement independently. Therefore, we assumed that the uneven distribution of glass eels observed in the habitat type experiments were the result of habitat preference. Given a choice of habitats in tank experiments, shortfin and longfin glass eels preferred habitats containing structure, and in particular, rocks/cobbles.     

Application of otolith microchemistry to estimate the migratory history of Japanese eel Anguilla japonica on the Sanriku Coast of Japan

The age and migratory history of the Japanese eel, Anguilla japonica Temminck & Schlegel, collected in Miyako Bay along the Sanriku coast of Japan, was examined using the otolith microstructure and analysis of strontium (Sr) and calcium (Ca) concentrations conducted with wavelength dispersive X‐ray spectrometry by an electron microprobe. The line analysis of Sr : Ca ratios along the life history transect of each otolith showed a peak (ca. 15–17 × 10^?3) which corresponded with the period of their leptocephalus and early glass eel stages in the ocean. The mean Sr : Ca ratios from the elver mark to the otolith edge indicated that there were eels with several general categories of migratory history, including sea eels that never entered freshwater (average Sr : Ca ratios, ≥6.0 × 10^?3), and others that entered freshwater for brief periods but returned to the estuary or bay. This evidence of the occurrence of sea eels in this northern area indicates that Japanese eels of the Sanriku coast do not necessarily migrate into freshwater rivers during recruitment as do glass eels at the beginning of their growth phase; even those that do enter freshwater may later return to the marine environment. Thus, anguillid eel migrations into freshwater are clearly not an obligatory migratory pathway, but rather a facultative catadromy with seawater or estuarine residents as an ecophenotype.     

Three-dimensional migratory behaviour of European silver eels (Anguilla anguilla) approaching a hydropower plant

The global population of European eel (Anguilla anguilla) is rapidly declining, and migration barriers in rivers are believed to be one of several key causes. While progress has been made in the development of bypass solutions, they are often constructed based on a limited knowledge of swimming behaviour. A bypass close to the stream bed is often recommended at fish passage facilities to accommodate downstream eel migration. The results of this recommendation are poorly studied, and the few studies that exist show varying bypass efficiencies. The current study used acoustic telemetry with depth sensors to explore the three-dimensional migratory behaviour of downstream-migrating silver eels. The eels were tracked as they approached a hydropower plant with a state-of-the-art angled bar rack and full-depth bypass. Downstream and upstream swimming differed in preferred vertical and lateral positions. During periods of local downstream movement, the density of observations was largest in the upper middle section, away from the river boundaries and in higher velocities. Conversely, when moving upstream, eels tended to avoid the upper layers of the middle part of the river, swimming closer to the riverbed and using the bank areas to a greater extent. Downstream-moving fish swam higher in the water column during night and in turbid conditions (high discharge). When approaching the impassable bar rack and the full-depth bypass, the eels searched most intensely but not exclusively along the bottom third of the rack, often exploring at new depths after changing direction. The impediment passage efficiency was 100% when both bypass solutions were considered. The study provides knowledge of the swimming behaviour of silver eels, which is relevant for the design of bypass solutions for eels at migration barriers.     

Occurrence of sea eels of <Emphasis Type="Italic">Anguilla japonica</Emphasis> along the Sanriku Coast of Japan

 The age and migratory history of the Japanese eel, Anguilla japonica, collected along the Sanriku Coast of Japan, were examined using otolith microstructure and analysis of strontium (Sr) and calcium (Ca) concentrations. The mean Sr : Ca ratios from the elver mark to the otolith edge indicated that there were eels with several general categories of migratory history, including sea eels that never entered freshwater and others which had entered freshwater for brief periods but returned to the estuary or bay. This first evidence of the occurrence of sea eels in this northern area indicates that Japanese eels of the Sanriku Coast do not necessarily migrate into freshwater rivers. Received: May 15, 2002 / Revised: August 4, 2002 / Accepted: August 15, 2002 Acknowledgments We thank Messrs. S. Yamane and K. Morita, and crews of the Otsuchi Marine Research Center, Ocean Research Institute, The University of Tokyo, for their assistance in collecting the eels. This work was supported in part by Grant-in-Aid No. 13760138 from the Ministry of Education, Culture, Sports, Science and Technology, Japan. Correspondence to:Takaomi Arai     

Biological control of invasive populations of crayfish: the European eel (<Emphasis Type="Italic">Anguilla anguilla</Emphasis>) as a predator of <Emphasis Type="Italic">Procambarus clarkii</Emphasis>

The red swamp crayfish, Procambarus clarkii, is a paradigmatic invader of freshwater systems. Several attempts have been made to mitigate its multiple impacts but none was successful. Among the different methods proposed, the use of the European eel (Anguilla anguilla) as an indigenous predator is promising but the available information about its predatory ability on crayfish is to date scanty. To fill this gap in knowledge, we ran three experiments in wetlands and irrigation ditches in Italy. The first experiment, in the laboratory, was aimed at quantifying the extent of predation by eels on crayfish, the second, in enclosures, the size classes of crayfish mainly preyed and the possible effect of the eels on P. clarkii behaviour, and the third, in the field, its ability to effectively reduce crayfish populations. Results showed that eels prey on small-sized or soft crayfish, attacking them from the back; an indirect effect was to reduce crayfish trophic activity, which in turn might increases crayfish mortality due to starvation and decreases impact on the community. However, as shown in the field, the use of eels should be appropriately calibrated to the context of application. Taken together, our results show that eels might be used as a complement to the traditional trapping method. However, additional studies are necessary to understand the adequate number of eels to be introduced and to develop appropriate methods for quantifying such effects.     

Anguilla rostrata glass eel migration and recruitment in the estuary and Gulf of St Lawrence

This study describes catches of Anguilla rostrata glass eels and associated oceanographic conditions in the St Lawrence Estuary and Gulf. Ichthyoplankton survey data suggest that they enter the Gulf primarily in May, migrate at the surface at night, and disperse broadly once they have passed Cabot Strait. They arrive in estuaries beginning at about mid-June and through the month of July. Migration extends west up to Québec City, in the freshwater zone of the St Lawrence Estuary, 1000 km west of Cabot Strait. Anguilla rostrata glass eels travel between Cabot Strait and receiving estuaries at a straight-line ground speed of c. 10–15 km day⁻¹. Catches of fish per unit effort in estuaries in the St Lawrence system are much lower than those reported for the Atlantic coast of Canada. Low abundance of A. rostrata glass eels in the St Lawrence system may be due to cold surface temperatures during the migration period which decrease swimming capacity, long distances from the spawning ground to Cabot Strait and from Cabot Strait to the destination waters (especially the St Lawrence River), complex circulation patterns, and hypoxic conditions in bottom waters of the Laurentian Channel and the St Lawrence Estuary.     

Critical swimming speed of yellow‐ and silver‐phase European eel (Anguilla anguilla,L.)

The prime objective of this study was to evaluate differences between the swimming performance of two distinct life stages of European eels. The critical swimming speed (U_crit) of 29 yellow‐ and 33 silver‐phase eels was evaluated in a swim tunnel. Silver‐phase eels showed a better swimming performance (U_crit = 0.66 ms^?1) than yellow individuals (U_crit = 0.43 ms^?1). Male and female silver eels reached an identical U_crit despite their different sizes, which may be a strategy to increase the synchronization of arrival at the spawning grounds.     

Migratory history and habitat use by New Zealand freshwater eels <Emphasis Type="Italic">Anguilla dieffenbachii</Emphasis> and <Emphasis Type="Italic">A. australis</Emphasis>, as revealed by otolith microchemistry

 The migratory history of Anguilla dieffenbachii and A. australis, collected from a coastal lake of New Zealand, was examined using analysis of strontium (Sr) and calcium (Ca) concentrations. Line analysis of Sr : Ca ratios along the life history transect of each otolith showed a peak (Ca. 16–20 × 10⁻³) between the core and elver mark, which corresponded to the period of their leptocephalus and early glass eel stages in the ocean. The mean Sr : Ca ratios from the elver mark to the otolith edge indicated that eels had different migratory histories, which included freshwater residency in some eels (average Sr : Ca ratios, 1.7 × 10⁻³–2.4 × 10⁻³) but not in others (average Sr : Ca ratios, 3.1 × 10⁻³–6.5 × 10⁻³). These findings suggest that New Zealand freshwater eels have a flexible migration strategy and an ability to adapt to various habitats and salinities. Received: November 25, 2002 / Revised: January 17, 2003 / Accepted: January 17, 2003     

Effects of body condition and water temperature on Anguilla anguilla glass eel migratory behavior

Glass eels arriving from the sea use alternative migratory tactics, leading either to the colonization of rivers or to an early settlement in marine or estuarine habitats. In the field, the migration may be environmentally affected by water temperature and the migratory behavior could be physiologically dependent on the body condition (energetic status). To investigate how these environmental and physiological effects on the migration are behaviorally mediated, we experimentally tested the effects of changes in water temperature and body condition on locomotor activity (upstream swimming) and salinity preference of Anguilla anguilla glass eels. Low water temperature reduced significantly both locomotor activity and preference for freshwater, in accordance with field data showing that low water temperatures hinder both the estuarine migration and river recruitment. Glass eels switched from a freshwater- towards a saltwater-preference as their body condition decreased, confirming that the energetic status may affect the migratory behavior. We suggest that, in the wild, this condition-dependent change in salinity preference of low body condition glass eels induces an early settlement in marine or estuarine habitats. Such a behavioral shift, stopping the energy expenditure linked to river-oriented migratory behavior, may be adaptive by limiting the probability of death due to exhaustion. Our results show that the glass eel migratory behavior, through locomotor activity and salinity preference, may be controlled by interacting physiological and environmental factors.     

Cost of transport and optimal swimming speed in farmed and wild European silver eels (Anguilla anguilla)

A swimming speed of 0.4 meters per second (m s(-1)) is the minimal speed for European female silver eels to reach the spawning sites in the Sargasso Sea in time. As silver eels cease feeding when they start their oceanic migration, the cost of transport (COT) should be minimised and the swimming speed optimised to attain the highest energetic efficiency. In this study, we have investigated the optimal swimming speed (U(opt)) of silver eels since U(opt) may be higher than the minimal swimming speed and is more likely to resemble the actual cruise speed. A variety of swimming tests were performed to compare endurance swimming between farmed eels and wild eels, both in freshwater and in seawater. The swimming tests were run with 101 silver female eels (60-96 cm, 400-1500 g) in 22 Blazka-type swim tunnels in a climatised room at 18 degrees C with running freshwater or seawater. Tests were run at 0.5-1.0 m s(-1) with increments of 0.1 m s(-1), and either 2 h or 12 h intervals. Remarkably, both tests revealed no changes in oxygen consumption (M O2) and COT over time. U(opt) values ranged between 0.61 and 0.68 m s(-1) (0.74-1.02 BL s(-1)) for the different groups and were thus 53-70% higher than the minimal speed. At U(opt), the COT was 37-50 mg O2 kg(-1) km(-1). These relatively very low values confirm our earlier observations. COT values in seawater were about 20% higher than in freshwater. Assuming that migrating female silver eels cruise at their U(opt), they will be able to cover the distance to the Sargasso Sea in 3-4 months, leaving ample time for final maturation and finding mates.     

Time lag of the response on the otolith strontium/calcium ratios of the Japanese eel, <Emphasis Type="Italic">Anguilla japonica</Emphasis> to changes in strontium/calcium ratios of ambient water

We conducted a laboratory experiment to validate the relationship between the otolith strontium/calcium (Sr/Ca) ratio of Japanese eels (Anguilla japonica) and water Sr/Ca ratio when the ratio in water was changed. A linear and additive mixed modeling approach was used to assess otolith Sr/Ca ratio for elver-juvenile Japanese eels when ambient water was changed from seawater to freshwater. There was a significant difference between otolith Sr/Ca ratios of eels reared in freshwater and in seawater (freshwater: 1.3–2.3; seawater: 7.0–7.8 mmol/mol). The response of otolith Sr/Ca ratios of eels was not detected until after 10 d and models suggested that it might not be completed until at least 30–60 d. This study indicated the detailed ability of otolith Sr/Ca ratio to be used as a proxy for reconstructing the individual environmental history of Japanese eels. These findings can provide some assurances for future otolith Sr/Ca studies of eels in this system or in other areas that have similar environmental conditions.     

Built to bite? Differences in cranial morphology and bite performance between narrow‐ and broad‐headed European glass eels

The presence of two phenotypes in a single species is a widespread phenomenon, also observed in European eel (Anguilla anguilla). This dimorphism has been related to dietary differences in the subadult elver and yellow eel stages, with broad‐heads generally feeding on harder and/or larger‐bodied prey items than narrow‐heads. Nevertheless, both broad‐ and narrow‐headed phenotypes can already be found among glass eels, the stage preceding the elver eel stage. As these glass eels are considered nonfeeding, we investigate here to what degree the observed variation in head width is reflected in variation in the musculoskeletal feeding system, as well as whether this reflects the same variation observed in the older, dimorphic yellow eels. Additionally, we investigate whether musculoskeletal differences between broad‐ and narrow‐headed glass eels have implications on their feeding performance and could thus impact prey preference when eels start feeding. Therefore, we compared the cranial musculoskeletal system of five broad‐ and narrow‐headed glass eels using 3D‐reconstructions and simulated the glass eel's bite force using the data of the muscle reconstructions. We found that the variation in the musculoskeletal system of glass eels indeed reflects that of the yellow eels. Broader heads were related to larger jaw muscles, responsible for mouth closure. Accordingly, broad‐heads could generate higher bite forces than narrow‐headed glass eels. In addition, broader heads were associated with higher coronoid processes and shorter hyomandibulae, beneficial for dealing with higher mechanical loadings and consequently, harder prey. We, thus, show that head width variation in glass eels is related to musculoskeletal differences which, in turn, can affect feeding performance. As such, differences in prey preference can already take place the moment the eels start feeding, potentially leading to the dimorphism observed in the elver and yellow eel stage.     

Behavioural responses of glass eels (Anguilla anguilla) towards amino acids

The behavioural responses of glass eels of Anguilla anguilla towards amino acids were investigated by binary choice experiments testing different concentrations of 14 L-amino acids in fresh water and salt water. Glass eels responded to solutions of individual amino acids down to a concentration of 10⁻⁸M, 10⁻⁹m. Media of different salinities influenced the responses. In freshwater, gln and thr were strongly attractive; asn, ala, met, glu and ile induced significant avoidance; gly elicited multimodal reactions depending on concentrations; his, lys, phe, val, leu and asp hardly influenced behaviour. In salt water, only gly, asn and lys significantly influenced the behaviour of the glass eels.     

Rhythmic swimming activity in Anguilla anguilla glass eels: Synchronisation to water current reversal under laboratory conditions

Synchronisation of swimming activity to water current reversal every 6.2 h was tested in the European glass eel (Anguilla anguilla L.). When presented with a change in water current direction, glass eels exhibited rhythmic patterns of activity with a period close to the tidal one. Glass eels began to swim with the current and then alternated between positive and negative rheotaxis after each change in the water current direction. Results are discussed in relation to the flood tidal transport theory. Following synchronisation to current reversal, glass eels subjected to constant conditions displayed a weak rhythmic activity suggesting that locomotor behaviour might, in the wild, synchronise to several environmental cues related to the tide. Results obtained with different densities also suggest that social cues might improve the synchronisation.     

Observations on the Nocturnal Activity and Feeding Behavior of Anguilla japonica Glass Eels Under Laboratory Conditions

Glass eels of the temperate anguillid species, Anguilla japonica, clearly showed a nocturnal activity rhythm under laboratory conditions. Light–dark cycle was a determinant factor affecting their photonegative behavior, nocturnal locomotor activity, and feeding behavior. Under natural light conditions, glass eels remained in shelters with little daytime feeding, but came out to forage during darkness. They moved and foraged actively in the following dark, and then their activity gradually declined possibly because of food satiation. They finally buried in the sand or stayed in tubes immediately after the lights came on. Under constant light, glass eels often came out of the shelters to forage in the lights but spent little time moving outside the shelters (e.g. swimming or crawling on the sand). Glass eels took shelter to avoid light and preferred tubes to sand for shelter possibly because tubes were much easier for them to take refuge in than sand. Feeding and locomotor activities of the glass eels were nocturnal and well synchronized. They appeared to depend on olfaction rather than vision to detect and capture prey in darkness. Feeding was the driving force for glass eels to come out of sand under constant light. However, in the dark, some glass eels swam or crept actively on sand even when they were fully fed. The lunar cycles of activity rhythms of glass eels that have been observed in some estuarine areas were not detected under these laboratory conditions.     

A comprehensive hypothesis on the migration of European glass eels (Anguilla anguilla)

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.