RNA/DNA ratios in American glass eels (Anguilla rostrata): evidence for latitudinal variation in physiological status and constraints to oceanic migration?

During their larval leptocephalus phase, newly hatched American eels undergo an extensive oceanic migration from the Sargasso Sea toward coastal and freshwater habitats. Their subsequent metamorphosis into glass eel is accompanied by drastic morphological and physiological changes preceding settlement over a wide geographic range. The main objective of this study was to compare RNA/DNA ratios and condition factor among glass eels in order to test the null hypothesis of no difference in physiological status and metabolic activity of glass eels at the outcome of their oceanic migration. This was achieved by analyzing glass eel samples collected at the mouth of 17 tributaries covering a latitudinal gradient across the species distribution range from Florida to Gaspésie (Québec). Our main observations were (i) a latitudinal increase in mean total length; (ii) a latitudinal variation in mean RNA/DNA ratios, which was best explained by a quadratic model reaching its minimum in the central range of sampling locations; and (iii) a latitudinal variation in Fulton's condition factor, which was best explained by a quadratic model reaching its maximum in the central range of sampling locations. Below we discuss the possible links between latitudinal variation in glass eel physiological status and variable energetic and environmental constraints to oceanic migration as a function of latitudinal distribution.     

Qualitative assessment of the diet of European eel larvae in the Sargasso Sea resolved by DNA barcoding

European eels (Anguilla anguilla) undertake spawning migrations of more than 5000 km from continental Europe and North Africa to frontal zones in the Sargasso Sea. Subsequently, the larval offspring are advected by large-scale eastward ocean currents towards continental waters. However, the Sargasso Sea is oligotrophic, with generally low plankton biomass, and the feeding biology of eel larvae has so far remained a mystery, hampering understanding of this peculiar life history. DNA barcoding of gut contents of 61 genetically identified A. anguilla larvae caught in the Sargasso Sea showed that even the smallest larvae feed on a striking variety of plankton organisms, and that gelatinous zooplankton is of fundamental dietary importance. Hence, the specific plankton composition seems essential for eel larval feeding and growth, suggesting a linkage between eel survival and regional plankton productivity. These novel insights into the prey of Atlantic eels may furthermore facilitate eel larval rearing in aquaculture, which ultimately may replace the unsustainable use of wild-caught glass eels.     

Oceanic fronts in the Sargasso Sea control the early life and drift of Atlantic eels

Anguillid freshwater eels show remarkable life histories. In the Atlantic, the European eel (Anguilla anguilla) and American eel (Anguilla rostrata) undertake extensive migrations to spawn in the oceanic Sargasso Sea, and subsequently the offspring drift to foraging areas in Europe and North America, first as leaf-like leptocephali larvae that later metamorphose into glass eels. Since recruitment of European and American glass eels has declined drastically during past decades, there is a strong demand for further understanding of the early, oceanic phase of their life cycle. Consequently, during a field expedition to the eel spawning sites in the Sargasso Sea, we carried out a wide range of dedicated bio-physical studies across areas of eel larval distribution. Our findings suggest a key role of oceanic frontal processes, retaining eel larvae within a zone of enhanced feeding conditions and steering their drift. The majority of the more westerly distributed American eel larvae are likely to follow a westerly/northerly drift route entrained in the Antilles/Florida Currents. European eel larvae are generally believed to initially follow the same route, but their more easterly distribution close to the eastward flowing Subtropical Counter Current indicates that these larvae could follow a shorter, eastward route towards the Azores and Europe. The findings emphasize the significance of oceanic physical–biological linkages in the life-cycle completion of Atlantic eels.     

Inter and intra-estuary variability in ingress, condition and settlement of the American eel Anguilla rostrata: implications for estimating and understanding recruitment

The objective of this study was to quantify spatial and temporal variability of anguillid glass eel ingress within and between adjacent watersheds in order to help illuminate the mechanisms moderating annual recruitment. Because single fixed locations are often used to assess annual recruitment, the intra-annual dynamics of ingress across multiple sites often remains unresolved. To address this question, plankton nets and eel collectors were deployed weekly to synoptically quantify early stage Anguilla rostrata abundance at 12 sites across two New Jersey estuaries over an ingress season. Numbers of early-stage glass eels collected at the inlet mouths were moderately variable within and between estuaries over time and showed evidence for weak lunar phase and water temperature correlations. The relative condition of glass eels, although highly variable, declined significantly over the ingress season and indicated a tendency for lower condition A. rostrata to colonize sites in the lower estuary. Accumulations of glass eels and early-stage elvers retrieved from collectors (one to >1500 A. rostrata per collector) at lower estuary sites were highly variable over time, producing only weak correlations between estuaries. By way of contrast, development into late-stage elvers, coupled with the large-scale colonization of up-river sites, was highly synchronized between and within estuaries and contingent on water temperatures reaching c. 10−12° C. Averaged over the ingress season, abundance estimates were remarkably consistent between paired sites across estuaries, indicating a low degree of interestuary variability. Within an estuary, however, abundance estimates varied considerably depending on location. These results and methodology have important implications for the planning and interpretation of early-stage anguillid eel surveys as well as the understanding of the dynamic nature of ingress and the spatial scales over which recruitment varies.     

Assessment of the environmental drivers of European glass eel (Anguilla anguilla) recruitment in transitional waters

The European eel (Anguilla anguilla) has undergone an unprecedented population decline since the 1980s, with current recruitment levels fluctuating from 3 to 15% of historical levels for the last 20 years. Monitoring of glass eels and elvers as 0?+?recruitment is an essential step in helping to understand the trend in recruitment and to better quantify the current recruitment time series. Two locations within the Shannon estuary on the west coast of Ireland were monitored for glass eel recruitment from January to April in 2017 and 2018. This study used a generalised linear mixed model to examine a range of environmental variables impacting on glass eel abundance in transitional waters. Results found that water temperature and moon phase were the most important variables. Tidal height and cloud cover also influenced the abundance of glass eels but to a lesser extent. This study found that focussing survey efforts on nights around the full moon when water temperatures exceed 5℃ will allow a catch which is representative of the population in an estuary. Glass eel monitoring needs a long-term sampling plan in order to account for annual fluctuations apparent in glass eel recruitment.

     

Sympatric spawning but allopatric distribution of Anguilla japonica and Anguilla marmorata: temperature- and oceanic current-dependent sieving

Anguilla japonica and Anguilla marmorata share overlapping spawning sites, similar drifting routes, and comparable larval durations. However, they exhibit allopatric geographical distributions in East Asia. To clarify this ecological discrepancy, glass eels from estuaries in Taiwan, the Philippines, Indonesia, and China were collected monthly, and the survival rate of A. marmorata under varying water salinities and temperatures was examined. The composition ratio of these 2 eel species showed a significant latitude cline, matching the 24 °C sea surface temperature isotherm in winter. Both species had opposing temperature preferences for recruitment. A. marmorata prefer high water temperatures and die at low water temperatures. In contrast, A. japonica can endure low water temperatures, but their recruitment is inhibited by high water temperatures. Thus, A. japonica glass eels, which mainly spawn in summer, are preferably recruited to Taiwan, China, Korea, and Japan by the Kuroshio and its branch waters in winter. Meanwhile, A. marmorata glass eels, which spawn throughout the year, are mostly screened out in East Asia in areas with low-temperature coastal waters in winter. During summer, the strong northward currents from the South China Sea and Changjiang River discharge markedly block the Kuroshio invasion and thus restrict the approach of A. marmorata glass eels to the coasts of China and Korea. The differences in the preferences of the recruitment temperature for glass eels combined with the availability of oceanic currents shape the real geographic distribution of Anguilla japonica and Anguilla marmorata, making them "temperate" and "tropical" eels, respectively.     

Influence of oceanic factors on Anguilla anguilla (L.) over the twentieth century in coastal habitats of the Skagerrak,southern Norway

The European eel (Anguilla anguilla L.) is distributed in coastal and inland habitats all over Europe, but spawns in the Sargasso Sea and is thus affected by both continental and oceanic factors. Since the 1980s a steady decline has been observed in the recruitment of glass eels to freshwater and in total eel landings. The eel is considered as critically endangered on the International Union for the Conservation of Nature and Natural Resources Red List of species. The Skagerrak beach seine survey from Norway constitutes the longest fishery-independent dataset on yellow/silver eels (starting in 1904). The Skagerrak coastal region receives larvae born in the Sargasso Sea spawning areas that have followed the Gulf Stream/North Atlantic Drift before they penetrate far into the North Sea. The Skagerrak coastal time series is therefore particularly valuable for exploring the impacts of oceanic factors on fluctuations in eel recruitment abundance. Analyses showed that Sargasso Sea surface temperature was negatively correlated with eel abundance, with a lag of 12 years revealing a cyclic and detrimental effect of high temperatures on the newly hatched larvae. The North Atlantic Oscillation index and inflow of North Atlantic water into the North Sea were negatively correlated with eel abundance, with a lag of 11 years. Increased currents towards the North Atlantic during high North Atlantic Oscillation years may send larvae into the subpolar gyre before they are ready to metamorphose and settle, resulting in low recruitment in the northern part of the distribution area for these years. The Skagerrak time series was compared with glass eel recruitment to freshwater in the Netherlands (Den Oever glass eel time series), and similar patterns were found revealing a cycle linked to changes in oceanic factors affecting glass eel recruitment. The recent decline of eels in the Skagerrak also coincided with previously documented shifts in environmental conditions of the North Sea ecosystem.     

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.     

Discrepancies between otoliths of larvae and juveniles of the American eel: is something fishy happening at metamorphosis?

The use of otoliths to interpret early life history in fishes depends upon the assumptions that otoliths record past events accurately and consistently and that records of events in otoliths are continuous. Both the number of growth microincrements ( I ), and the radii ( R ,μm) of otoliths of American eel Anguilla rostrata , leptocephali increased linearly and highly significantly with leptocephalus body length ( L , mm), as expected on the above assumptions ( I , =2·29 L , − 5·75 and R , =1·05 L , + 12·02, r ²,=0·938 and 0·931, n , =20). In contrast, the number of increments and the radii of the leptocephalus growth zones of otoliths of glass-phase American eels were not related to body length, and they were lower than predicted by the relationships developed for leptocephali. Thus, otoliths of American eels apparently violate one or both assumptions. Possibly, the margin of the otolith is resorbed during metamorphosis from leptocephalus to glass eel, perhaps as part of calcium metabolism as skeletal elements are being formed.     

The early life history of the tropical eel Anguilla marmorata (Quoy & Gaimard, 1824) from four Pacific estuaries, as revealed from otolith microstructural analysis

Anguilla marmorata glass eels or elvers were collected separately during anadromous migration from four Pacific estuaries: Hamuta, Poso, Shuang Hsi and Tanshui. The total length at arrival in these estuaries was (mean ± standard error) (51.50 ± 0.90) (51.80 ± 0.90) (46.95 ± 0.84) and (47.33 ± 0.80) mm, respectively. The sagittal otolith microstructure, increment patterns and daily age were examined by scanning electron microscope. Based on the number of increments of presumed daily deposition, the overall mean age at arrival in the estuaries was estimated to be about 3–4 months, with an estimated period of 73–86 days for the leptocephalus stage. Two zones, i.e. the leptocephalus growth zone (L) and the metamorphosis growth zone (M) were recognizable in the otolith cross section. The increment width of L and M varied from the otolith's centre to its margin, reflecting different growth rates. The spawning grounds of these eels are presumably not far from the estuary. Their locations are discussed.     

Head shape dimorphism in European glass eels (Anguilla anguilla)

The life cycle of the European eel (Anguilla anguilla) remained a mystery until the 20th century, when Schmidt discovered that the Sargasso Sea was its spawning area. However, many aspects of the eel's life cycle remain poorly understood. Among these is the bimodal distribution in head shape, with broad- and narrowheaded phenotypes reported in the yellow eel stage. Although this has been linked to dietary preferences of the yellow eels, very little is known about why, how and when this dimorphism arises during their ontogeny. To determine whether this dimorphism indeed appears in relation to trophic niche segregation, we examined head shape variation at an earlier ontogenetic stage, the glass eel stage, as at this stage eels are considered to be non-feeding. Head shape was studied in a large dataset, containing glass eels captured from the Yser river mouth, the Leopold Canal (Belgium) and from the rivers Severn, Trent and Parret (UK), by both taking measurements (head width/head length) and using an outline analysis. Our results show that there is already considerable variation in broadness and bluntness of the head at the glass eel stage. In most cases, equal support for a unimodal and bimodal head shape distribution is found, whereas some cases support head shape bimodality in glass eels, suggesting that glass eel head shape might be shifting from a unimodal to a bimodal distribution. This, in combination with the observation that variation in head width/head length ratios in non-feeding glass eels shows a similar range as in feeding yellow eels, indicates that head shape in European eel might be at least partially determined through other mechanisms than trophic segregation.     

The Seasonal Occurrence of Glass Eels (Anguilla Anguilla L.) in the Rio Minho between 1991 and 1993 (North of Portugal)

Glass eels were sampled monthly at two stations in the Rio Minho. The experimental fishery was carried out in the estuary between November 1991 and July 1992 and at an upstream site between December 1991 and March 1993. The yield differed at each station, especially in summer and between the official seasons of the experimental glass eel fishery. Early pigmentation stages were more frequent in the estuary and the maximum lengths and weights were observed in autumn and winter. The official statistics of the professional glass eel fishery are briefly mentioned.     

Long-term numerical changes and regulation in a river stock of European eel Anguilla anguilla

1. The objective was to explore the regulatory mechanisms underlying the numerical dynamics of a river stock of European eel Anguilla anguilla based on a purely observational study. We used a 21-year data set on the Rio Esva (Asturias, northwestern Spain) stock of eels, encompassing eel density at an estuary site and at nine sites distributed among three tributaries selected to differ in the distance from the tidal limit and fishery data on glass eels from the nearby Rio Nalón estuary.
2. The abundance of glass eels and estuarine and in-stream densities declined simultaneously over a 15-year period. However, following a historical minimum in the abundance of glass eels around the year 2000, the density continued to decline in the estuary whereas density in the streams began an increase that has continued to the present.
3. In-stream cohort mortality rate ( Z ) varied from 0.0016 day⁻¹ to only 0.0003 day⁻¹ (on an annual basis from 0.57 to 0.099 year⁻¹). The negative effects of intra-cohort and inter-cohort densities on mortality rate emphasized the responses of the riverine stock to compensatory density-dependence. As a consequence, the number of eels surviving to commence the silvering stage was similar in magnitude in the strongest and weakest cohorts.
4. These results suggest that, even if the low abundance of glass eels persists, in-stream density may increase towards a stable value regulated by in-stream density-dependence.     

Compilation of Japanese fisheries statistics for the Japanese eel, <Emphasis Type="Italic">Anguilla japonica</Emphasis>, since 1894: a historical dataset for stock assessment

Fishery sustainability and the extinction risk of the Japanese eel, Anguilla japonica, are of global concern. The landings of the Japanese eel in Japan comprise a large part of the landings in East Asia. This study provides a compiled dataset of the annual fisheries statistics of the Japanese eel in Japan for stock assessment. The Japanese government has been recording Japanese eel statistics annually since 1984 in five series of annual reports by conducting systematic questionnaire surveys of fisheries managers and associations; however, most of these data are stored in analog format. The key variables in the dataset include the harvest weight of eels, the harvest weight and number of seeds for aquaculture, the number of eels stocked, and the number of management entities engaged in the eel fishery. The levels of spatial aggregation of the variables include the site (river and lake), prefecture, inland and coastal waters, and total in Japan. We also incorporated location data (latitude and longitude) of the site and prefecture into the dataset. Eel harvest includes primarily yellow eels (late juvenile stage) and silver eels (mature stage). Seed harvest in inland waters includes glass eels (intermediate stage between leptocephalus and elver) and elvers (early juvenile stage). Seed harvest from coastal waters comprises glass eels. This dataset provides information to assess long-term trends in the Japanese eel population.     

A review of glass eel migratory behaviour,sampling techniques and abundance estimates in estuaries: implications for assessing recruitment,local production and exploitation

Recruitment of the three northern hemisphere eel species (European eel Anguilla anguilla, American eel Anguilla rostrata and Japanese eel Anguilla japonica) has reduced significantly over the past thirty-five years. The stock of the European eel is described as being outside safe biological limits, with urgent action required by European Union Member States to assist recovery of the panmictic stock. Stock recruitment models and estimates of silver eel output from a river catchment are strongly influenced by the degree of certainty in estimating key population parameters of each life history stage. Therefore, management decisions aimed at enhancing eel populations rely on sound scientific evidence, based upon a fundamental understanding of the complex anguillid eel life cycle. This review paper focuses on the estuarine entry phase of the eel life cycle and synthesises the current scientific knowledge with regard to glass eel migratory behaviour, sampling methods and abundance estimates within estuaries. Although the behavioural and environmental processes modulating glass eel migration patterns are reasonably well understood, site specific factors play a significant role in determining fine scale distribution patterns at an individual estuary level. Given the large resource commitment required to adequately sample this key life history stage, behavioural studies of migration patterns on a local scale are crucially important to aid the design of robust sampling programmes aimed at quantifying seasonal abundance and annual recruitment.     

Long-term effects of hydropower installations and associated river regulation on River Shannon eel populations: mitigation and management

The Shannon, Ireland’s largest river, is used for hydroelectricity generation since 1929. Subsequently, the Electricity Supply Board assumed responsibility for management of its eel stocks, due to the impact of the hydro-dam on recruitment to the commercial fishery. In order to negate a decline in juvenile recruitment resulting from the installation of hydroelectric facilities, management was focused on stocking lakes with elvers and fingerling eels. These were trapped at the hydropower facilities and in estuarine tributaries during their up-stream migrations. Due to the decline of natural recruitment in more recent times, attempts have also been made to develop an estuarine glass eel fishery. Stock levels are then monitored through annual surveys of the population trends of juvenile (glass eel, elver), growing phase (yellow eel) and downstream migrating pre-spawners (silver eels). Survey results and fishery management programmes are reviewed in this article. In addition to the long-term effects the hydroelectric facilities have had on the stock levels, there is also an annual effect on the migratory patterns of downstream migratory silver eels. In the lower reaches of the river system flow rates are regulated by the hydroelectric stations. We review previous work that had highlighted the importance of flow in determining the timing of the silver eels migrations, and assess the relationship between flow and migration in more detail through the use of hydroacoustic and telemetric studies. Current research on seaward migrating silver eel populations, suggests that spawner escapement rates can most effectively be increased by trapping migrating eels at fishing weirs located up-stream of the power station and transporting them towards the estuary. Guest editors: R. L. Welcomme & G. Marmulla Hydropower, Flood Control and Water Abstraction: Implications for Fish and Fisheries     

长江口日本鳗鲡降海产卵洄游群体的银化指标研究

以长江口降海生殖洄游途中的日本鳗鲡为研究材料, 对繁殖群体银化过程中体色、年龄、全长、体重及银化指标等特征作了分析。结果显示, 依据“体侧黑色素沉积是否蔓延至胸鳍下缘”这一体色标准, 可将日本鳗鲡洄游群体分为银化前期和银化期二个发育期。日本鳗鲡银化过程中, 体型、胸鳍长、眼径、性腺指数呈递增趋势, 消化道指数则呈递减趋势。主成分分析亦表明, 银化期较银化前期的眼径和胸鳍增大, 还具有较高的性腺指数和较低的消化道饱满度。由于日本鳗鲡的产卵场位于3000 km之遥的马里亚纳海沟, 这些形态的修正可能是为了适应即将起始的长距离降海生殖洄游。根据银化指标所建立的判别方程, 对雌、雄群体银化前期和银化期的判别分析均取得了95%以上的成功率。研究认为, 长江口日本鳗鲡银化期的野外快速判别, 亦可采用“眼径指数大于5”作为识别依据。     

长江靖江段沿岸日本鳗鲡丰度的时间格局及生物学研究

于2002-2017年在长江靖江段沿岸对日本鳗鲡资源量进行长期调查,对其丰度的时间格局特征及环境影响因素等进行了分析.结果 显示,在近16年519次调查中,采集到日本鳗鲡137尾,平均年龄(1.8±1.0)龄,平均全长和体质量分别为(31.1±9.8) cm和(57.64±91.94)g,89.1％的个体未性成熟,94...     

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.     

Recruitment abundance estimation: Role of glass eel (Anguilla anguilla L.) response to light

Most fish populations are declining worldwide and their management would benefit from a better estimation of recruitment. In glass eels, field studies suggest that estuarine migratory glass eels are sensitive enough to light to change their vertical location according to factors such as water turbidity and/or moon brightness. The response of glass eel (Anguilla anguilla L.) to light was tested in the laboratory using boxes where fish could choose between a lit and an unlit side. Responses were quantified as the proportion of glass eels remaining in the unlit chamber. Decreasing light levels were used and tested on different “age” glass eels (“age” in days since capture). In addition, measures of light at different depths of the water column were carried out in the Adour estuary (43°30′ N, 1°30′ W). The glass eel light avoidance level was lower in non-pigmented glass eel (less than 10 ^− 10 W cm ^− 2), than in pigmented ones (10 ⁻⁹-10 ^− 8 W cm ^− 2). These results and field data on the measurement of light energy in the water column of Adour estuary are compared with previously published data on the estuarine migration of glass eel.