Salinity-linked growth in anguillid eels and the paradox of temperate-zone catadromy

Temperate-zone anguillid eels use both saline (marine or brackish) and fresh waters during their continental phase, but use of fresh waters is paradoxical because on average these fishes grow more rapidly in saline than in fresh waters. Based on data from anguillid eels whose habitat-residency histories had been determined by Sr:Ca otolithometry, superiority of growth rates in saline water is much greater in American eels Anguilla rostrata in north-eastern North America (mean saline:fresh growth rate ratio 2·07) than in European Anguilla anguilla , Japanese Anguilla japonica and shortfinned Anguilla australis eels (range of mean ratios 1·12–1·14). Data from A. rostrata in the Hudson Estuary, U.S.A., and Prince Edward Island, Canada, were used to test adaptive explanations of catadromous migrations. The hypothesis that lower mortality in fresh water offsets faster growth in saline water was not supported because loss (mortality + emigration ) rates did not vary between saline and fresh zones of the Hudson Estuary. Hypotheses that anguillid eels move to fresh water to escape from larger anguillid eels in saline water or to evaluate habitat quality were not supported by size and age distributions. Catadromy in temperate-zone anguillid eels increases the diversity of occupied habitats and therefore lowers fitness variance caused by environmental fluctuations. Catadromy in temperate-zone anguillid eels could be due to natural selection for maximum geometric mean fitness which is sensitive to fitness variance. Temperate-zone catadromy might also be maladaptive, at least in local areas, due to shifts over time in selective pressures or to inability of panmictic genetic systems to adapt to local conditions.     

How does salinity influence habitat selection and growth in juvenile American eels Anguilla rostrata?

The influence of salinity on habitat selection and growth in juvenile American eels Anguilla rostrata captured in four rivers across eastern Canada was assessed in controlled experiments in 2011 and 2012. Glass eels were first categorized according to their salinity preferences towards fresh (FW), salt (SW) or brackish water (BW) and the growth rate of each group of elvers was subsequently monitored in controlled FW and BW environments for 7 months. Most glass eels (78–89%) did not make a choice, i.e. they remained in BW. Salinity preferences were not influenced by body condition, although a possible role of pigmentation could not be ruled out. Glass eels that did make a choice displayed a similar preference for FW (60–75%) regardless of their geographic origin but glass eels from the St Lawrence Estuary displayed a significantly higher locomotor activity than those from other regions. Neither the salinity preferences showed by glass eels in the first experiment nor the rearing salinities appeared to have much influence on growth during the experiments. Elvers from Nova Scotia, however, reached a significantly higher mass than those from the St Lawrence Estuary thus supporting the hypothesis of genetically (or epigenetically) based differences for growth between A. rostrata from different origins. These results provide important ecological knowledge for the sustained exploitation and conservation of this threatened species.     

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.     

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.     

Plasma non-esterified fatty acid profiles and 17β-oestradiol levels of juvenile immature and maturing adult American eels in the St Lawrence River

American eels Anguilla rostrata , collected from three distinct locations along the St Lawrence River (Lake St Lawrence, Quebec City and Kamouraska), were thought to consist entirely of out-migrating adults at Quebec City and Kamouraska and of both resident and migrants at Lake St Lawrence. The mean plasma levels of 17β-oestradiol closely paralleled the trend observed with gonadosomatic index (I_G) and oocyte development. The highest levels of 17βoestradiol and I_G were observed in Quebec City and Kamouraska, while the levels observed in the resident eels from Lake St Lawrence were very low. Generally, the concentration of total plasma non-esterified fatty acids (NEFA) appeared to follow the trend of 17β-oestradiol and I_G and increased with sexual maturity. However, the most sexually mature group, eels from Kamouraska, had the lowest total plasma NEFA levels. The reason for the drop is not clear at present. In addition, a number of shifts in the plasma NEFA profiles of American eels did occur during their migration in a downstream direction, with several fatty acids either increasing or decreasing in relative abundance. The ratio of n3: n6 fatty acids declined with increasing sexual maturity as most n3 fatty acids became less abundant. Although the correlation between gonadal development and the relative concentration of individual fatty acids was not always clear, both arachidonic acid (20: 4n6) and docosahexaenoic acid (22: 6n3) were significantly higher in the plasma of out-migrating adults than in resident yellow phase eels.     

Anguilla rostrata glass eel ingress into two, U.S. east coast estuaries: patterns, processes and implications for adult abundance

A time series of American eel Anguilla rostrata glass eel abundance, timing and size from Little Egg Inlet, New Jersey (16 years) and Beaufort Inlet, North Carolina (18 years) was used to provide a better understanding of ingress patterns at two, U.S. east coast estuaries. There was no evidence of synchronous declines in abundance between the two locations; however, at the Little Egg Inlet site, glass eels arrived later in the season and at significantly smaller sizes over the duration of the series. One significant linkage between sites was revealed: abundance was positively correlated with winter precipitation. Precipitation differed between sites annually and was correlated with El Niño at Beaufort Inlet and, to a lesser extent, the North Atlantic Oscillation at Little Egg Inlet. It is hypothesized that glass eels may use freshwater signals to enhance recruitment to local estuaries, thus influencing year-class strength, yet the relationship between year-class strength and adult abundance remains unresolved.     

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.     

Shark Predation on Migrating Adult American Eels (Anguilla rostrata) in the Gulf of St. Lawrence

In an attempt to document the migratory pathways and the environmental conditions encountered by American eels during their oceanic migration to the Sargasso Sea, we tagged eight silver eels with miniature satellite pop-up tags during their migration from the St. Lawrence River in Québec, Canada. Surprisingly, of the seven tags that successfully transmitted archived data, six were ingested by warm-gutted predators, as observed by a sudden increase in water temperature. Gut temperatures were in the range of 20 to 25°C—too cold for marine mammals but within the range of endothermic fish. In order to identify the eel predators, we compared their vertical migratory behavior with those of satellite-tagged porbeagle shark and bluefin tuna, the only endothermic fishes occurring non-marginally in the Gulf of St. Lawrence. We accurately distinguished between tuna and shark by using the behavioral criteria generated by comparing the diving behavior of these two species with those of our unknown predators. Depth profile characteristics of most eel predators more closely resembled those of sharks than those of tuna. During the first days following tagging, all eels remained in surface waters and did not exhibit diel vertical migrations. Three eels were eaten at this time. Two eels exhibited inverse diel vertical migrations (at surface during the day) during several days prior to predation. Four eels were eaten during daytime, whereas the two night-predation events occurred at full moon. Although tagging itself may contribute to increasing the eel''s susceptibility to predation, we discuss evidence suggesting that predation of silver-stage American eels by porbeagle sharks may represent a significant source of mortality inside the Gulf of St. Lawrence and raises the possibility that eels may represent a reliable, predictable food resource for porbeagle sharks.     

Discrimination of New Zealand stream waters by glass eels of Anguilla australis and Anguilla dieffenbachii

This study tested the hypotheses, using glass eels of longfin eels Anguilla dieffenbachii and shortfin eels Anguilla australis migrating into fresh waters, (1) that both species prefer water from their river of collection to well water, (2) that shortfin eels prefer lowland, pastoral stream water to mainstem river water, (3) that longfin eels are attracted to both waters but do not prefer one to the other, and (4) both prefer water scented with geosmin, a widely occurring metabolite of bacteria and algae, to well water. Glass eels of both species from a river on the west coast of South Island, New Zealand, and shortfin glass eels from an east coast river significantly preferred water from their river of capture to well water. Two to three times as many eels chose their own river water as chose well water. Longfin eels were rare in the east coast river. Shortfin glass eels from the two rivers chose lowland stream water to mainstem river water about two to one in three experiments with different pairs of waters to which they had no prior exposure. Longfin glass eels significantly chose mainstem river water over lowland water in one pair but showed no preference when presented with a different pair. Reactions to solutions of geosmin at concentrations of 10^–5-10^–7 mg 1^–1 were inconclusive, with geosmin being preferred significantly, by shortfin eels, in only one experiment. The interspecific differences in discrimination of natural waters demonstrated in this study, with shortfin eels preferring lowland waters and longfin eels more indifferent to water types, are in broad agreement with both the distribution of adults and observations on their habitat preferences.     

Temporal and spatial distribution of the ichthyotoxic dinoflagellate Gyrodinium aureolum Hulburt in the St Lawrence, Canada

The occurrence of the potentially ichthyotoxic dinoflagellateGyrodinium aureolum Hulburt is reported for the first time inthe Lower Estuary and Gulf of St Lawrence. Taxonomic identificationusing an iramunochemical tagging method suggests a close taxonomicproximity between the species found in the St Lawrence and G.aureolumpresent in European waters. In 1993, G.aureolum was observedalong the entire coast of Quebec and in offshore waters of theGulf of St Lawrence. At the coastal stations, G.aureolum showedlarge week-to-week variations, but higher densities were observedduring late August and early September. Bloom concentrations(10⁶ cells 1^–1) were only observed in the GaspéCurrent at Mont-Louis, on the north coast of the Gaspd peninsula.In the central part of the Gulf, G.aureolum was concentratedin the upper 5 m of the stations directly influenced by thefreshwater run-off of the St Lawrence Estuary. Results fromprincipal component analysis indicate that G.aureolum favorsenvironments with high nitrogen recycling activity.     

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.     

Validation of daily otolith increments in glass-phase American eels Anguilla rostrata (Lesueur) during estuarine residency

Prior to making inferences from otoliths about the residence time and growth rate of glass-phase anguillid eels Anguilla in estuaries, it is necessary to validate the deposition rate of microincrements in the otoliths. Glass-phase American eels Anguilla rostrata (Lesueur), which had been captured near the mouth of an estuary in Maine, USA, prior to freshwater exposure, deposited increments at a daily rate at ambient temperature and salinity in a field and laboratory study. The regression for glass eels not possessing a transition ring was: I=0.976(D-1)+0.434, where I is the number of otolith increments distal to a fluorescent mark placed on the otolith at the beginning of the experiment, and D is the number of days in the experiment, which ranged from 7 to 49. The slope was not significantly different than 1. Unexpectedly, many glass eels deposited the transition ring during the experiment, although this ring had previously been thought to mark entry into fresh water. The regression for these glass eels was: I=0.961(D-1)-3.880, and the slope was not significantly different than 1. The negative intercept suggests that approximately 4 days were lost from the otolith record during deposition of the ring. This study demonstrated daily deposition of increments prior to freshwater exposure and demonstrated that deposition of the transition ring is not linked to freshwater entry.     

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.     

The exotic American eel in Taiwan: ecological implications

Six American eels Anguilla rostrata were identified in the Kaoping River of south-western Taiwan by sequencing the mitochondrial cytochrome b gene ( mtCyt-b ). This was the first record of an exotic eel species in the natural waters of Taiwan. The exotic eels may have been imported from North America at the elver stage for aquaculture and escaped from culture ponds.     

Spring migration of Atlantic mackerel,Scomber scombrus,in relation to water temperature through Cabot Strait (Gulf of St. Lawrence)

Synopsis Sea surface temperatures across Cabot Strait (Gulf of St. Lawrence) ranged from 6 to 9°C on June 3, 1989, but only from 3 to 5°C on June 2, 1990. Periods of peak commercial landings of mackerel in eastern Cape Breton Island extended from May 22 to June 3 in 1989, and from May 28 to June 2 in 1990. In late May 1990, Atlantic mackerel were captured with a purse-seiner during exploratory fishing in Cabot Strait; commercial quantities were caught in water as cold as 2.8°C. The presence of mackerel in water a full 4°C colder than its reported lower tolerance limit indicates that the development of the 7°C isotherm is not a requirement for the vernal appearing of mackerel. The overlap of the periods of peak commercial landings between 1989 and 1990, despite: marked differences in warming chronology, suggests that the movements of mackerel are not as closely linked to water temperature as previously reported. The fish's thermal preferences could be subordinate to their reproductive requirements at this stage of their spawning migration.     

Differential effects of origin and salinity rearing conditions on growth of glass eels of the American eel Anguilla rostrata: implications for stocking programmes

In this study, growth patterns were monitored in controlled fresh and brackish water (BW) conditions for 7 months during Anguilla rostrata glass eel and elver stages. Null hypotheses tested were that there is no significant difference in growth between glass eels (1) collected from two geographic regions typified by different sex ratios, (2) reared in fresh and BW and (3) due to origin × salinity interactions. It was found that young A. rostrata from Mira River (MR, Cape Breton, Nova Scotia, Canada, an area where both males and females occur) grew faster than those from Grande-Rivière-Blanche (Québec, Canada, an area where population are highly skewed towards females; 99–100%). Anguilla rostrata from both origins also grew faster in BW, although there was a trend for origin × salinity interactions whereby this effect was more pronounced for fish from the MR. The results support the hypothesis that salinity can influence growth patterns, as possibly can quantitative genetic differences between A. rostrata glass eels from different origins. Possible explanations for these patterns and potential consequences for sex determination and translocation programmes are discussed.     

Yellow-stage American eel movements determined by microtagging and acoustic telemetry in the St Jean River watershed, Gaspé, Quebec, Canada

The hypothesis that a part of the yellow American eel Anguilla rostrata sub-population of the St Jean River in eastern Quebec feeds in the brackish environment during summer and returns to the river to overwinter was tested. Three years of microtagging and the acoustic tagging and tracking of 40 American eels demonstrated that a part of the downstream migrants exploited the estuary as a summer feeding area. Upstream movement of some microtagged American eels provided support for the hypothesis that a part of those American eels returned to the river to overwinter. In addition to the demonstration of amphidromous behaviour of yellow eels, the study revealed that American eels in the estuary were active at night but homed to specific daytime resting sites.     

Effectiveness of a strobe light eel exclusion scheme

In the laboratory and field, American eels (Anguilla rostrata) showed a strong avoidance of white strobe light. Eels avoided low light intensities for all strobe flash frequencies tested, and showed no behavioural adaptation to the light source over a prolonged time period. A strobe light barrier was 65–92% effective in repelling upstream migrating eels at Saunders GS on the St. Lawrence River. All size classes of eels were repelled, but effectiveness appeared to be reduced for smaller eels.     

Anguilla japonica is already magnetosensitive at the glass eel phase

Magnetosensitivity of the Japanese eel Anguilla japonica at the glass eel phase (newly metamorphosed juveniles) was examined by conditioning and electrocardiography. The glass eels were conditioned to an imposed magnetic field of 192 473 nT parallel to the fish body placed along the earth's west‐east axis. After 10 to 40 conditioning runs, all the glass eels exhibited a significant conditioned response ( i.e . slowing of the heart beat) to a 192 473 nT magnetic field and even to a 12 663 nT magnetic field that combined with the geomagnetic field (32 524 nT) at the laboratory and produced a resultant magnetic field of 21° easterly. These results indicate that glass eels have high magnetosensitivity and probably acquire geomagnetic information early in life. It is hypothesized that silver‐phase adult eels find their way back to the oceanic spawning ground by reversing the geomagnetic direction that had been detected and 'memorized' during the glass eel phase when migrating from the open ocean towards the continental shelf and coastal waters.