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.     

APPLIED ISSUES: Size‐dependent mortality of migratory silver eels at a hydropower plant,and implications for escapement to the sea

1. The European eel population has decreased drastically during recent decades, and new EU‐legislation calls for measures to change this negative trend. This decline has been attributed to a number of factors, including habitat fragmentation by structural barriers that prevent eels moving between freshwater and the sea. The success of downstream migrating adult silver eels migrating past a hydroelectric plant (HEP) in Sweden was examined by radio‐telemetry, and the results were considered in a historical context by analysing catch data from the river for 1957–2006. 2. The choice of routes and passage success were quantified for three treatment groups and one control group of silver eels. The first treatment, the reservoir group (n = 50), was released into the reservoir upstream of the HEP, and these fish could proceed downstream by passing through the HEP (20 mm rack and turbines) or by entering the spill gates into the former channel, bypassing the HEP. The second treatment group (inside rack, n = 15) was released downstream of the 20‐mm rack and had to pass through the turbines to continue migration to the sea. The third treatment group consisted of dead radio‐tagged eels (n = 6) that were released into the turbines to study the extent of drifting by dead individuals. Finally, the control group (n = 50) was released downstream of the HEP to test for effects of confounding factors. 3. Most live individuals displayed migratory behaviour and continued to proceed downstream after release. Only 8% of the fish released in the reservoir or downstream of the HEP (control) did not migrate. The probability of reaching the next HEP, 24 km further downstream, was high for the control group (96%) and the reservoir‐released individuals that passed the HEP via the spill gates and the former channel (83%). Survival was low and size‐dependent for the individuals that passed the turbines (40%) and even lower for the individuals that had to pass through the rack and the turbines (26%). The overall passage success for eels released in the reservoir was 30%, including both routes. 4. Annual catch data from 1957 to 2006 showed that the number of eels in the River Ätran has decreased. Despite this decrease, escapement biomass has remained unchanged, because of the fact that the mean size of eels has doubled. Passage data from 2007 show that changes in size and abundance have resulted in a reduction of relative escapement to the sea to values that are 21–24% of what they were in 1957–66. However, this low level of escapement could potentially be rectified if appropriate measures facilitating HEP passage are successfully implemented, since the potential escapement biomass in the river, owing to the large size of the eels, has changed little since the 1950s.     

Success of a low‐sloping rack for improving downstream passage of silver eels at a hydroelectric plant

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Loss of European silver eel passing a hydropower station

The aim of this study was to assess escapement success of silver eels, Anguilla anguilla (L.), in a lowland river while passing a reservoir and a hydropower station. It was hypothesized that passage success would be lowest at the hydropower station and that survival and migration speed would be highest in the free‐flowing river section upstream the reservoir. Forty‐five female silver eels 56–86 cm in length were tagged with acoustic transmitters and released in November 2006. Their migration was monitored via automatic listening stations (ALS) in various sections of the river, covering a total migration distance of 64 km. Survival and progression rate of downstream migration was highest in the upstream river section and significantly lower in the reservoir. The eels apparently had trouble finding their way past the turbines and spent between 1.5 and 35 h in the forebay. The results show that within the study period, only 23% of the tagged eels reached the tidal limit, mainly due to difficulties in passing the hydropower dam. With such high loss‐rates, the escapement goals set in the management plan cannot be achieved.     

Passage survival of European and American eels at Francis and propeller turbines

We present the magnitude of losses of European eel Anguilla anguilla and American eel A. rostrata in passage through propeller and Francis turbines at hydroelectric projects. Survival and injury rates and types were turbine type related. Overall, eel survival was higher (mean ± 90% CI = 95.1 ± 5.3%,) and injury rate lower (12.5 ± 10.5) at Francis than propeller turbines (survival = 80.7 ± 6.4%; injury rate = 25.7 ± 7.9%). The common injury type at Francis turbines was bruises and at propeller turbines was severance. Blade shape and thickness of the leading edge of the blades (rounded, thick buckets of Francis turbines v. flatter, sharper edged blades in propeller turbines); eel entry routes into the turbines; their flexible, cylindrical body shape and orientation probably contributed to these differences. Relationship between survival and injury and turbine characteristics was turbine specific. For Francis turbines, one negative correlation (r = −0.986, P < 0.01) between survival and runner speed was found and two positive correlations between injury rates and fish length (r = 0.740, P < 0.10) and number of blades (r = 0.835, P < 0.05) were noted for propeller turbines. Several severely injured eels remained active 48 h after turbine passage suggesting caution is warranted when using telemetric movement for estimating eel survival. We conclude there is a need to (a) better understand travel paths and approach orientation of eels through turbines; (b) determine where only eel passage is of concern at hydropower plants that have both turbine types and therefore preferential operation of Francis turbines may be considered; (c) inform hydropower plant operators where turbine replacement is being considered and downstream eel passage is of concern that replacement by Francis or bulb turbines may prove beneficial for eel passage.     

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.     

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.     

Variations in the migratory history of the tropical catadromous eels Anguilla bicolor bicolor and A. bicolor pacifica in south-east Asian waters

Fish movements between aquatic habitats of different salinity ranges (fresh, estuarine, marine) by the tropical catadromous eels Anguilla bicolor bicolor and A. bicolor pacifica were examined by analysing the otolith strontium and calcium concentrations of yellow (immature) and silver (mature) stage eels collected in south-east Asian (Indonesia, Malaysia and Vietnam) waters. The ratios suggest that all migratory-type eels, including freshwater, brackish water and marine residents, pass the river mouth. However, the habitat preference was different among the sites (countries). In Indonesia and Vietnam, most A. bicolor bicolor and A. bicolor pacifica were either marine or brackish water residents in this study. Alternatively, most A. bicolor bicolor were freshwater residents in Malaysia; such a typical catadromous migration pattern in these eels has not been found in previous studies. The wide range of otolith Sr:Ca in both subspecies indicates that the habitat use of these tropical eels was opportunistic among fresh, brackish and marine waters during their growth phases following recruitment to coastal areas. The geographical variability of migratory histories suggests that habitat use might be determined by the inter and intraspecific competition and environmental conditions at each site.     

Optimizing the mass marking of fish with alizarin red S: an example with glass eels

Fish marking is an essential tool for fisheries management, especially for evaluating the stocking of endangered fish species to support conservation and sustainable use of fish stocks. Batch marking of young European eels Anguilla anguilla (L.) prior to stocking is recommended as the benefits of stocking for the spawning stock can be evaluated by recapturing marked fish over time, therefore mass marking of young eels with substances such as alizarin red S (ARS) is becoming increasingly important. To improve the marking method and reduce marking costs when immersing glass eels in an ARS solution, eight laboratory experiments under varying conditions (e.g., temperature, ARS concentration, immersion time, osmotic induction, fish density) and with ARS from different suppliers were carried out. The results show that optimal marking of glass eels can be carried out in the field or during transport by putting approximately 50 g of glass eels per liter in 150 mg L⁻¹ ARS solution for 3 h at 10–15°C. Lower concentrations did not result in reliable marking. Water temperatures of 5°C and below can have a stunning effect on the eels and increase mortality significantly, regardless of the concentration of ARS. Glass eel densities below 50 g L⁻¹ in the marking bath increase marking costs unnecessarily, while a higher density of 100 g L⁻¹ resulted in significantly higher mortality and lower marking success. A somewhat more difficult but less expensive alternative is to bathe the fish in a saline solution of 1% (10 PSU) of 80 mg L⁻¹ ARS for 3 h at 10°C. Costs can also be significantly reduced by choice of supplier for ARS, but care should be taken as the quality of the powder appears to vary (mean percentage of sufficiently marked eels ranged from 59% to 91% among suppliers in the present study) and can lead to marking failure. The optimal marking conditions can help ensure that stocked glass eels can be reliably identified in future studies to assess stocking benefits while reducing costs.     

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.     

Electrolyte changes of serum and muscle,and related mortalities in maturingAnguilla rostrata migrating down the St. Lawrence Estuary (Canada)

This study was made to investigate changes in serum and muscle ion concentrations and related mortalities in maturingAnguilla rostrata migrating down the St. Lawrence Estuary. Mortalities take place in the freshwater portion of the St. Lawrence. Electrolyte concentrations of moribund eels taken in freshwater were compared to those of freshwater and salt water controls. Moribund eels had a much lower serum osmolality (270 mOsm/kg) than the controls (328 and 358 mOsm/kg). This resulted from low sodium (125 mEq/l) and particularly low chloride (69 mEq/l) contents in the moribund eels compared to the freshwater controls (153 and 117 mEq/l) and the salt water controls (179 and 137 mEq/l). There was also a general decrease in muscle ion concentrations in moribund eels though the percentage water was similar to that of the freshwater controls (64.0 and 63.7%). The changes measured between the freshwater controls and the salt water controls in nature are similar to those measured onAnguilla anguilla in laboratory. These results suggest that mortalities are related to failure by some of the maturing eels to maintain their mineral balance in freshwater. Hypothesis is made that maturing eels migrating long distances in freshwater or retarded by physical or chemical barriers, start to excrete sodium and chloride under hormonal control before they have reached brackish water. In the conditions that prevail in the St. Lawrence Estuary, this results in mineral unbalance and possibly in mortalities.     

Effects of dietary carbohydrates and temperatures on slow growing juvenile eels Anguilla anguilla

Synopsis The influence of the level of carbohydrate in a purified diet on the growth of slow-growing Anguilla anguilla maintained at different temperatures was examined. The average weight increase of eels maintained at temperatures of 25° or 27° C and fed a diet containing 20% or 30% glucose, was significantly higher than the mean weight increase of eels maintained at the same temperature but fed a diet containing soluble corn starch, at the same percentages. There was no significant difference in the mean weight of slow growing eels fed 10%, 20% and 30% soluble corn starch or 10% glucose. There was no significant difference in protein content (wet weight) among the experimental groups. However, elvers that were fed a high percentage of carbohydrates (glucose or starch) and maintained at 27° C had a higher percentage of lipids in body weight compared with the other experimental groups.     

Validation of annulus in otolith and estimation of growth rate for Japanese eel <Emphasis Type="Italic">Anguilla japonica</Emphasis> in tropical southern Taiwan

The age of Japanese eels (Anguilla japonica) is often estimated from otoliths, but this method has not been fully validated, particularly in tropical areas where the annulus in otolith is considered to be less distinct than in temperate areas. To validate the annuli in Japanese eel otoliths from southern Taiwan, known-age (2 year-old) cultured eels from an eel farm and wild eels from Kao-Ping River were collected. It was found that 26 out of 31 cultured eels (83.9%) showed two clear annuli and the remained 5 eels showed either one or three annuli. The mean (± SD) age of the cultured eels was 1.97 ± 0.4 years. Meanwhile, a clear peak in the mean monthly marginal increment ratio of the otolith in wild yellow and silver eels occurred once a year during winter (November to March). The annual deposition of presumed annuli in otoliths of Japanese eel was validated and the age and growth rate estimation for Japanese eels in the tropical southern Taiwan is deemed feasible. The growth rate of cultured eels was significantly faster than that of wild eels, but it did not differ significantly between sexes for wild silver, yellow or cultured eels. The von Bertalanffy Growth Function parameters (K, and t ₀ ) of the wild eels were estimated as 0.114 ± 0.028 year⁻¹, 1178 ± 171 mm and −0.8 ± 0.2 years, respectively.     

A description of rodlet cells from the alimentary canal of Anguilla anguilla and their relationship with parasitic helminths

Rodlet cells in intestinal epithelia of infected and uninfected European eels Anguilla anguilla from brackish and fresh water were studied by light and electron microscopy. Deropristis inflata (Trematoda) was found in eels from brackish water, whereas eels from fresh water were infected with Acanthocephalus clavula (Acanthocephala). In a comparison between uninfected and infected eels from brackish water, a higher number of rodlet cells was recorded in the intestinal epithelia of infected fish. Evidence is presented that rodlet cells secrete their contents in a holocrine manner into the lumen of the eel intestine. The occurrence of organelles within the mature rodlet cell was rare. ? 1998 The Fisheries Society of the British Isles     

Relationship between elver recruitment and changes in the sex ratio of silver eels Anguilla anguilla L. migrating from Lough Neagh, Northern Ireland

Between 1932–1947 and from 1960 onwards, elvers have been trapped near the mouth of the River Bann, Northern Ireland, and released into Lough Neagh. Each period of elver transport has been followed by a marked increase in the proportion of male silver eels migrating from the lough. Catches of silver eels were sampled on several nights each year from 1965–1974, and the lengths of a total of 20358 eels measured showed a progressive increase in the percentage of male eels from 9.3-86.0 % during this period. Various reasons for this change were examined. The different ages at which male and female eels migrate to the sea was not important. There was no evidence to support the hypothesis that male elvers normally remain in estuarine conditions, and their transport to the lough was therefore unnatural. An increasing fishing effort for yellow eels, such as occurred following the introduction of trawling in 1960, would favour males since small eels were returned to the lough. It was not thought, however, that this was a major cause of the change in sex ratio. Instead, elver transport appeared to be directly implicated, possibly by the overstocking of Lough Neagh, and the phenotypic determination of progressively more male eels, but the evidence for this suggestion was inconclusive.     

Diel periodicity and chronology of upstream migration in yellow-phase American eels (<Emphasis Type="Italic">Anguilla rostrata</Emphasis>)

Yellow-phase American eel (Anguilla rostrata) upstream migration is temporally punctuated, yet migration chronology within diel time periods is not well-understood. This study examined diel periodicity, chronology, and total length (TL) of six multi-day, high-count (285–1,868 eels) passage events of upstream migrant yellow-phase American eels at the Millville Dam eel ladder, lower Shenandoah River, West Virginia during 2011–2014. We categorized passage by diel periods (vespertine, nocturnal, matutinal, diurnal) and season (spring, summer, late summer/early fall, fall). We depicted passage counts as time-series histograms and used time-series spectral analysis (Fast Fourier Transformation) to identify cyclical patterns and diel periodicity of upstream migration. We created histograms to examine movement patterns within diel periods for each passage event and fit normal mixture models (2–9 mixtures) to describe multiple peaks of passage counts. Periodicity of movements for each passage event followed a 24-h activity cycle with mostly nocturnal movement. Multimodal models were supported by the data; most modes represented nocturnal movements, but modes at or near the transition between twilight and night were also common. We used mixed-model methodology to examine relationships among TL, diel period, and season. An additive-effects model of diel period + season was the best approximating model. A decreasing trend of mean TL occurred across diel movement periods, with the highest mean TL occurring during fall relative to similar mean values of TL for spring, summer, and late summer/early fall. This study increased our understanding of yellow-phase American eels by demonstrating the non-random nature of their upstream migration.     

An Examination of Utilizing External Measures to Identify Sexually Maturing Female American Eels, Anguilla Rostrata, in the St. Lawrence River

It is well established that Anguillid eels undergo a complex suite of morphological and physiological changes during their transformation from resident, yellow-phase juveniles to actively migrating silver-phase eels. While it has been shown that some morphological measures can be used successfully to identify sexually maturing European eels, Anguilla anguilla, as well as Australian short fin, Anguilla australis, and long fin, Anguilla dieffenbachii eels, this relationship has never been quantitatively assessed for American eels, Anguilla rostrata. American eels of varying sexual development were collected from three locations on the St. Lawrence River: Lake St. Lawrence, Quebec City and Kamouraska. Sexual development of each eel was assessed with gonadosomatic index (GSI), oocyte diameter and degree of oocyte development. Morphological measures of total length, weight, head width, pectoral fin length and vertical and horizontal eye diameters were obtained from each fish. We used this data to test two hypotheses: (i) resident yellow phase eels, suspected migrants and known migrants are morphologically indistinguishable; and (ii) if differences exist, they cannot be used to reliably predict gonadal development or migratory status. Univariate analysis (ANOVA and ANCOVA) indicated that there were highly significant differences in all of the measured parameters and thus we were able to reject the first hypothesis. However, we failed to reject the second hypothesis as the high degree of overlap between groups eliminated the ability of any single measure to differentiate between resident and migratory eels. A multivariate discriminant model was developed that could classify only 72–80% of the eels correctly based on their morphological characters. While morphological measures may have some potential as a rapid, cost-effective method of pre-screening individual eels, morphological measures should not be considered a definitive indicator of sexual maturity or migratory status for female American eels in the Upper St. Lawrence River.     

Age and growth of migrating tropical eels,Anguilla celebesensis and Anguilla marmorata

The age and growth of migrating tropical eels, Anguilla celebesensis and Anguilla marmorata from central Sulawesi, Indonesia, were examined. Migrating eels (63 A. celebesensis and 38 A. marmorata ) were obtained from weirs near the Poso Lake outlet and non‐migrating eels (35 A. celebesensis and 119 A. marmorata ) were captured by baited hooks, eel pots, scoop net and electro‐fishing in the Poso River system, Laa River system, Baluga River, Tongku River and Padapu River from February 2009 to October 2010. In both species, the proportion of eels with opaque otolith edges showed a single peak in July, suggesting that one annulus (a pair of translucent and opaque zones) was formed each year in their otoliths. Mean ± s.d . and range of total length (L _T) and age was 785·2 ± 114·9 (585–1083) mm and 7·5 ± 1·6 (5–11) years in migrating female A. celebesensis and 1132·2 ± 173·7 (800–1630) mm and 11·6 ± 3·3 (7–23) years in A. marmorata . The age of migrating female eels was negatively correlated with annual growth rate, 100·7 ± 17·2 (68·1–145·0) mm year^?1 in A. celebesensis and 97·9 ± 19·3 (66·6–131·6) mm year^?1 in A. marmorata , but there was no significant correlation between the L _T and annual growth rate in either species. The annual growth rates of these female tropical eels were typically higher than those of temperate anguillid species, suggesting a latitudinal cline in growth rate in the genus Anguilla reflecting the environmental conditions of their growth habitat.     

Decline in non-native freshwater eels in Japan: ecology and future perspectives

The occurrence, distribution, and biological characteristics of non-native freshwater eels were analyzed using 5524 eels collected from 16 sites in Japan between 1997 and 2005. Three hundred seventy-four fishes (6.8%) were identified as non-native European eels, Anguilla anguilla, while the remainder (93.2%) were native Japanese eels, A. japonica. The European eel was found at 7 sites (44%), including 3 rivers, 2 freshwater lakes, one brackish lake, and one sea bay, suggesting a wide rage of habitat use. This variability of habitat use was also evidenced by the otolith microchemistry, which showed that they had lived in not only freshwater but also in seawater habitats. The sites with European eel were localized within the vicinity of southern Japan where a number of these eels were cultivated in the early 1970’s, suggesting that some had escaped from the culture ponds or were released intentionally into nearby natural waters. The large body size (mean total length: 803 mm), pigmented skin, enlarged eyes, and relatively matured gonads (mean gonad somatic index: 1.9) found in non-native European eels indicated that most had metamorphosed into the migratory silver phase, suggesting their ability to initiate spawning migration. However, the proportion of European eels in Mikawa Bay in 1997 was more than 12%, which decreased markedly to less than 2% after 2001, corresponding to the recent decline in import of European glass eels for aquaculture. This suggests that the population of European eels will decrease in Japanese waters in the future.