Metabolic response under different salinity and temperature conditions for glass eel Anguilla japonica

Diadromous fish often enter freshwater directly from seawater via fish ladders or channels built in estuarine dams. The oxygen consumption rates (OCR) of glass eel, Anguilla japonica, were determined using an automatic intermittent flow respirometer under various salinity and temperature regimes to physiologically explain this direct movement. The endogenous rhythm of the OCR in wild glass eels, freshly collected from estuaries, was nearly synchronous with the tidal pattern at the estuarine collection site. When the salinity was changed from 20 psu (12°C) at a constant temperature to that of freshwater, the OCR of the glass eels decreased by 21.6±7.0% (mean ± SD) (P<0.05), showing a dampened rhythm for about 48 h. After this period of impediment, the glass eels resumed normal metabolic activity. Direct migration from seawater to freshwater under constant temperature would result in a severe physiological stress for these glass eels for about two days. When the glass eels were exposed to a cyclic change in water temperature of 2°C 26 h⁻¹, as they encounter in estuaries, and then were introduced to freshwater abruptly, the OCR rhythm corresponded to the cyclic changes in water temperature after exposure to freshwater. Under these conditions, the mean OCR of the glass eels had a small difference before and after exposure to freshwater. These data explained how glass eels can directly move from sea water into the freshwater without any apparent metabolic stress in the estuaries showing cyclic change in water temperature (Δt=2°C).     

Changes in water and ion transport in isolated intestines of the eel during salt adaptation and migration

When Japanese cultured eels, Anguilla japonica, were transferred from freshwater to sea-water, their body weight decreased by 15% on the second day and then recovered gradually to the original weight within a week. Serum sodium content increased by 58% on the second day, and thereafter decreased to a constant level. Isolated intestines from these eels were bathed on both sides, mucosal and serosal, by identical Ringer solution. The water and sodium transort was very active from mucosa to serosa in such isolated intestines of sea-water-adapted eels. This active movement was particularly dominant during the week after transferring the eel to seawater, and a peak was reached on the fifth day. The molar ratio of water to sodium in the fluid moving across the intestinal wall was higher in sea-water-adapted eels than in freshwater eels. The isolated intestines taken from silver eels during their catadromous migration, but still in the river, showed a larger amount of water transport and a higher molar ratio of water to sodium than intestines from yellow sedentary eels. These adaptive properties are a prerequisite for the marine life of the eel.     

Rhythmic motor activity responses of the California fiddler crab Uca crenulata to artificial light conditions

Uca crenulata, the California species of fiddler crab, was exposed to artificial light conditions to separate the influence of the light cycle from that of the tidal input on its rhythmic motor activity. Under both constant light and light-dark cycles, rhythmic activity was demonstrated in only 50% of the experimental crabs; the activity of the remaining 50% was random. Individuals exposed to constant light conditions after having been subjected to 24 h light-dark cycles demonstrated no significant difference in period length of their rhythmic activity from crabs investigated in constant light immediately after field collection. The mean period did not differ significantly from the tidal period of 24.8 h, but the variation was considerable. In artificial light-dark cycles, the activity rhythms were usually masked but, in some cases, synchronized. The results indicate that U. crenulata has an endogenous rhythm with a period close to the tidal cycle which may be synchronized by light as well as by tidal cues. The display of this endogenous rhythm, however, is poor.     

Rhythmic activity responses of the fiddler crab Uca crenulata to artificial tides and artificial light

The fiddler crab Uca crenulata, from California (USA), was exposed to artificial tides in order to differentiate between the influence of a tidal cycle from that of a light-dark cycle on its locomotor activity. Most crabs could be well synchronized by tides, but the activity patterns of other crabs was merely exogenously reinforced. Under constant conditions, after tidal treatment, crabs showed bimodal or unimodal activity patterns. When a 24 h light-dark cycle and a 12.35 h tidal cycle acted as concurrent stimuli, their effectiveness in synchronizing the crabs' activity was variable, depending on the undividual. It is likely that crabs which tend to exhibit a unimodal activity pattern are more sensitive to a light stimulus, whereas crabs with a bimodal activity pattern respond preferably to the tides. Thus, responses of endogenous activity of U. crenulata to tides and light-dark cycles are similar, suggesting that the basic oscillarory mechanism for circadian and tidal activity may be the same.     

Changes in intestinal absorption and renal excretion of water during adaptation to sea-water in the Japanese eel

The rate of sea-water ingestion in Japanese cultured eels, Anguilla japonica, adapted to sea-water for one week, was 89±9.5 ml/kg/day; approximately 75% of the ingested water was absorbed by the intestine. Similar rates of ingestion and absorption of water were also found with sea-water-adapted rainbow trout, Salmo gairdnerii, and two species of marine teleosts, Stephanolepis cirrhifer and Goniistius zonatus. When cultured eels were transferred from freshwater to sea-water the amount of ingested and absorbed water inoreased gradually to a maximum on the 5^th day and then decreased to a constant level. The urine flow and glomerular filtration rate were found to be high in freshwater cultured eels, and both reduced markedly within 6 h after transferring the eel to sea-water. The reabsorption of water by the renal tubules was very small in both the freshwater and sea-water-adapted (6 h) eels. On the other hand, in the eels adapted to sea-water for 10 days the rate of urine flow was also very slight but the glomerular filtration rate recovered to that of the freshwater eels. Thus, water reabsorption by the tubules increased to a large degree. The results seem to indicate the existence of two distinct phases in the renal function during sea-water adaptation of cultured eels.     

Biology of the conger eel Conger oceanicus in the Mid-Atlantic Bight

In the Mid-Atlantic Bight, conger eels (Conger oceanicus) occur from the coastal portions of estuaries to the edge of the continental shelf. In deeper waters they occupy burrows of the tilefish (Lopholatilus chamaeleonticeps). Between 1972 and 1974 we examined the stomachs and intestines of conger eels from inshore New Jersey (USA) waters (n=35, with a total length: TL of 21 to 49 cm) and between 1980 and 1983 offshore (n=295, 50 to 125 cm TL)_collections. Eels from both areas fed primarily on decapod crustaceans and fish. The specific identity of prey items within these groups generally differed from inshore to offshore areas, probably reflecting the differences in prey availability. Foods of specimens collected offshore varied with size: smaller eels (<80 cm TL) fed most heavily on decapod crustaceans, whereas larger eels (>80 cm) consumed more fishes. The presence of some nocturnally active prey items in the gut, primarily the eel Lepophidium cervinum, suggests that conger eels are nocturnal feeders. This is supported by in situ observations that conger eels are present in some tilefish burrows during the day and are presumably out of burrows and foraging at night.     

Social synchronization of the activity rhythms of honeybees within a colony

Colonies and isolated bees of the Cape honeybee, Apis mellifera capensis Esch., were observed for evidence of circadian rhythmicity under constant conditions. It was found that colonies develop free-running activity rhythms in self-selected light-dark cycles, which are slightly shorter than 24 h. The periods of the activity rhythms of individual isolated bees were longer than 24 h in self-selected light-dark and constant light, while they were shorter than 24 h in constant darkness. A greater variability in period was found in the isolated bees than in the colonies. When the rhythms of colonies and individual bees from these colonies were measured simultaneously, the activities of the isolated bees drifted with respect to that of the colonies, their period being either longer or shorter than that of their own colony. After 12 days of isolation of individual bees from their colony, all coincidence between the phases of the two rhythms was lost. We conclude that the periods of common activity and common rest of the bees within a colony result from a mutual (social) synchronization of the rhythms of the individual bees.     

Relative contribution of migratory type on the reproduction of migrating silver eels, <Emphasis Type="Italic">Anguilla japonica</Emphasis>, collected off Shikoku Island,Japan

In order to understand the reproductive contribution among migratory types in the Japanese eel, Anguilla japonica, otolith strontium (Sr) and calcium (Ca) concentrations by X-ray electron microprobe analysis were examined for 37 silver eels collected in Kii Channel off Shikoku Island during the spawning migration season. The wide range of otolith Sr:Ca ratios indicated that the habitat use of A. japonica was not obligatory but facultative among fresh, brackish and marine waters during their growth phases after recruitment to the coastal areas as glass eels. Three migratory types, which were categorized as river eels, estuarine eels and sea eels were found. The estuarine eels were dominant (59%), followed by sea eels (22%) and river eels (19%). The low proportion of river eels from the spawning migration season suggested that the estuarine and sea eels inhabiting the nearby coastal areas might make a larger reproductive contribution to the next generation in this area.     

Occurrence of marine resident tropical eel Anguilla bicolor bicolor in Indonesia

In order to understand the migratory history and habitat use of the tropical anguillid eels Anguilla celebesensis, A. marmorata, and A. bicolor bicolor, the otolith strontium (Sr) and calcium (Ca) concentrations were examined for eels collected in Indonesian waters. In A. bicolor bicolor collected in a lagoon, the change in Sr:Ca ratios outside the high Sr:Ca core generally indicated two patterns of habitat residence: (1) constant living in either brackish or sea waters with no freshwater life (25%) and (2) habitat shifts from fresh water to brackish or sea waters (75%). No A. bicolor bicolor had a general life history as a freshwater resident. A. celebesensis and A. marmorata from the uppermost freshwater lake showed freshwater life history patterns. The wide range of otolith Sr:Ca ratios in A. bicolor bicolor indicated that the habitat use of this tropical eel was facultative among fresh, brackish, and marine waters during the growth phase after recruitment to coastal areas similar to that for temperate eels. Thus, the migration of anguillid eels into fresh water is clearly not an obligatory.     

Movement patterns of American eels (Anguilla rostrata) between salt- and freshwater in a coastal watershed, based on otolith microchemistry

Otolith strontium:calcium ratios were used to trace lifetime movements of American eels (Anguilla rostrata) captured in salt-water bays and adjoining freshwater ponds in Prince Edward Island, Canada. Eels were classified into migratory contingents based on their movement patterns. A pond with a pool-and-weir salmonid fishway and a pond drained by a low-gradient channel contained eels that had entered freshwater at all ages, but a pond with a 2.2 m vertical spillway contained only eels that had entered freshwater in the elver year. Salt-water residents were the dominant migratory contingent in salt-water bays (85% of 39), which overturns the paradigm of obligate catadromy for this species. Freshwater residency was the sole pattern found in the pond with the vertical spillway (100% of 12) and the majority contingent in the pond with the low-gradient channel (54% of 24). Inter-habitat shifting was the dominant migratory contingent in eels sampled from the pond with the pool-and-weir fishway (85% of 20). Resident eels were established in salt- and freshwater habitats by the year after their arrival in continental waters. Eels that shifted between habitats increased their rate of inter-habitat shifting with age. The high degree of plasticity in habitat use found in this study is consistent with worldwide Anguillid patterns as revealed by Sr:Ca.An erratum to this article can be found at     

Anguilliform larvae collected off North Carolina

The distinctive larval stage of eels (leptocephalus) facilitates dispersal through prolonged life in the open ocean. Leptocephali are abundant and diverse off North Carolina, yet data on distributions and biology are lacking. The water column (from surface to 1,293 m) was sampled in or near the Gulf Stream off Cape Hatteras, Cape Lookout, and Cape Fear, North Carolina during summer through fall of 1999–2005, and leptocephali were collected by neuston net, plankton net, Tucker trawl, and dip net. Additional samples were collected nearly monthly from a transect across southern Onslow Bay, North Carolina (from surface to 91 m) from April 2000 to December 2001 by bongo and neuston nets, Methot frame trawl, and Tucker trawl. Overall, 584 tows were completed, and 224 of these yielded larval eels. The 1,295 eel leptocephali collected (combining all methods and areas) represented at least 63 species (nine families). Thirteen species were not known previously from the area. Dominant families for all areas were Congridae (44% of individuals, 11 species), Ophichthidae (30% of individuals, 27 species), and Muraenidae (22% of individuals, ten species). Nine taxa accounted for 70% of the overall leptocephalus catches (in order of decreasing abundance): Paraconger caudilimbatus (Poey), Gymnothorax ocellatus Agassiz complex, Ariosoma balearicum (Delaroche), Ophichthus gomesii (Castelnau), Callechelys muraena Jordan and Evermann, Letharchus aliculatus McCosker, Rhynchoconger flavus (Goode and Bean), Ophichthus cruentifer (Goode and Bean), Rhynchoconger gracilior (Ginsburg). The top three species represented 52% of the total eel larvae collected. Most leptocephali were collected at night (79%) and at depths > 45 m. Eighty percent of the eels collected in discrete depth Tucker trawls at night ranged from mean depths of 59–353 m. A substantial number (38% of discrete depth sample total) of larval eels were also collected at the surface (neuston net) at night. Daytime leptocephalus distributions were less clear partly due to low catches and lower Tucker trawl sampling effort. While net avoidance may account for some of the low daytime catches, an alternative explanation is that many species of larval eels occur during the day at depths > 350 m. Larvae of 21 taxa of typically shallow water eels were collected at depths > 350 m, but additional discrete depth diel sampling is needed to resolve leptocephalus vertical distributions. The North Carolina adult eel fauna (estuary to at least 2,000 m) consists of 51 species, 41% of which were represented in these collections. Many species of leptocephali collected are not yet known to have juveniles or adults established in the South Atlantic Bight or north of Cape Hatteras. Despite Gulf Stream transport and a prolonged larval stage, many of these eel leptocephali may not contribute to their respective populations.     

Contrasts between spawning times of <Emphasis Type="Italic">Anguilla</Emphasis> species estimated from larval sampling at sea and from otolith analysis of recruiting glass eels

This study reviewed literature on spawning times for three north temperate species of anguillid eels estimated by sampling for small leptocephali (larvae) at sea and for several temperate and tropical species by back-calculating from putative daily ages derived from otolith increment analysis of glass eels that recruited to coastal waters. Estimates from otoliths of European eels, Anguilla anguilla, American eels, Anguilla rostrata, and Japanese eels, Anguilla japonica, imply much more protracted spawning seasons than are indicated by sampling at sea during various times of year. European eels are inferred to spawn year-round from otolith analysis, but the smallest, recently hatched leptocephali are found only in late winter and spring. From otoliths, the spawning times of these three species are all estimated to occur much later in the year than when small leptocephali are found at sea, indicating that ages appear to be underestimated. For these and other temperate and tropical eels, there are inconsistencies in assigned ages among various studies, which are most extreme for the European eel. This species has the longest larval migration and often has an opaque zone in the glass eels’ otoliths where it is difficult to discern growth increments. These inconsistencies suggest that interpretation of otolith growth increments is incorrect at least in some studies, and the apparently consistent mismatch between otolith and sea-sampling studies suggests that increments may not always be formed at some period during the unusual early life history of anguillids. Because daily increments may be formed in eels during most of their early life history, future research is needed to determine the cause of the mismatch of glass eel aging studies and the apparent spawning times of eels offshore. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.

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Anguilliform fishes and sea kraits: neglected predators in coral-reef ecosystems

Despite intensive sampling efforts in coral reefs, densities and species richness of anguilliform fishes (eels) are difficult to quantify because these fishes evade classical sampling methods such as underwater visual census and rotenone poisoning. An alternative method revealed that in New Caledonia, eels are far more abundant and diverse than previously suspected. We analysed the stomach contents of two species of sea snakes that feed on eels (Laticauda laticaudata and L. saintgironsi). This technique is feasible because the snakes return to land to digest their prey, and (since they swallow their prey whole) undigested food items are identifiable. The snakes’ diet consisted almost entirely (99.6%) of eels and included 14 species previously unrecorded from the area. Very large populations of snakes occur in the study area (e.g. at least 1,500 individuals on a small coral islet). The snakes capture approximately 36,000 eels (972 kg) per year, suggesting that eels and snakes play key roles in the functioning of this reef ecosystem.     

The effect of mating on female sexual behavior of Teleogryllus commodus Walker

Summary Virgin Teleogryllus commodus females show rhythmic locomotor activity in light-dark (LD) and constant light (LL) conditions. A taped 3-h calling song stimulates them into further walking, and they lay few eggs. Successful mating suppresses or reduces circadian-controlled walking, but in LL-conditions the free-run still persists. Mated females no longer respond to the 3-h calling song. The number of eggs laid within 24–72 h after insemination increases, and the three effects of mating occur independently of one another. The possible physiological basis of these changes and their ecological and behavioral significance under field conditions are discussed.     

Molt cycle alterations in behavior,feeding and diel rhythms of a decapod crustacean,the spiny lobster Panulirus argus

The crustacean molt cycle manifests extensive behavioral changes in addition to physiological and integumentary modifications. The paucity of quantitative studies led us to characterize molt stage dependent alterations in rhythmic locomotor, feeding, and agonistic behavior of subadult spiny lobster Panulirus argus held grouped and solitarily in simulated natural conditions. Non-disruptively determined molt stages were defined as proportions of intermolt duration. Significant nocturnal rhythmicity persisted through the full molt cycle, though daily form varied as a function of activity level and molt stage. A distinct early evening peak signaled initiation of foraging and walking behavior. Thereafter, rhythm amplitude either declined progressively (low activity: unimodal pattern), remained high (high activity: nocturnal plateau), or exhibited a secondary early morning peak (intermediate activity: bimodal and polymodal patterns). Activity ceased at or immediately prior to sunrise. Ecdysis was predominantly nocturnal, probably reflecting species specific spatiotemporal movement patterns and social behavior. A selective advantage of nocturnal ecdysis was postulated concerning avoidance of cannibalism and diurnal predators. Locomotor activity and feeding rates were not equivalent through the molt cycle, though both peaked in stages B₂-C₁. Locomotor activity remained high in metecdysis, decreased sharply at proecdysis initiation (D₀), reaching lowest levels in D₁-D₃, then increased in D₃ through B₁. Activity dropped steeply at ecdysis, though lobsters were capable of intense and coordinated activity. Feeding decreased slowly through metecdysis after the B₂-C₁ maximum, then increased temporarily in C₄ and D₀, indicating heightened feeding motivation. This contrasts with the locomotor activity decrease at proecdysis. Food consumption declined rapidly in D₁ and D₂ and ceased at the D₂-D₃ transition. Feeding resumed in B₁ or B₂, intensifying to maximum in late B₂. Feeding remained relatively constant within stages, whereas locomotor activity varied greatly, though both correlated with metabolic needs. Grouped and solitary lobsters displayed similar patterns of foraging and walking, equivalent to those of locomotor activity and feeding of solitary individuals. Frequency of agonistic interactions (not aggression per se) remained relatively constant through the cycle, peaking in metecdysis, though the highest relative proportion occurred near ecdysis. Lobsters then were submissive and avoided physical contact with conspecifics. Clearly, locomotor activity, feeding, and social behavior of P. argus are not simply determined. Indeed, behavior is distinctly phase coordinated with varying metabolic requirements dependent on the proximity to ecdysis and ecological pressures.     

Exploratory observations on microhabitat selection within the intertidal zone by the Chinese mudskipper fish Periophthalmus cantonensis

The amphibious Chinese mudskipper fish Periophthalmus cantonensis liver in complex, variable, tropical intertidal mangrove forest and mudlfat habitats. Within these macrohabitats, an array of microhabitats are available to this species which spends most of its time out of water. It is subjected to, and tolerates, wide ranges of salinity and temperature. It is primarily diurnally active and uses dark, usually water-filled burrows in the mud as refuges and breeding places. Individuals of P. cantonensis were captured at night from intertidal mudflats of mangrove forests in the northeastern New Territories, Hong Kong in 1971 and 1972. Using choice experiments in the laboratory, we determined behavioural preferences of this fish for five combinations of abiotic environmental variables relevant to its selections of microhabitats. P. cantonensis usually preferred being out of water, resting on damp mud, to being in direct contact with water of any salinity. In salinity-choice experiments it showed no preference for any particular salinity, but avoided exposure to fresh water. Offered choices between various water temperatures it displayed a strong preference for higher temperatures (30° to 35°C), even though these temperatures were close to those known to cause distress or death. In light-dark choice experiments in very shallow water P. cantonensis preferred darkness to light. Experiments combining watertemperature differences with light-dark choices showed that these two factors interact in complex ways. We discuss implications of these results for understanding of behavioural selection of microhabitats by this fish.     

三丁基锡和菲对鳗鲡的毒性效应（Toxic Effects of Tributyltin and Phenanthrene on Eels）

采用腹腔注射方式(每周1次,共5次)将生活于淡水或海水中的日本鳗鲡(Anguilla japonica)暴露于三丁基锡氯化物(TBTCl)和菲(Phe),以研究这两种污染物对鳗鲡的毒性效应以及该效应与鱼体所处环境盐度的关系.鳗鲡暴露于TBTCl或Phe1w后,其肝脏和脾脏的组织结构就已发生明显病变,且病变程度随着暴露时间的延长而加剧,其中海水鳗鲡比淡水鳗鲡表现更为严重;但肝体指数(HSI)和性腺组织结构均无明显变化.暴露于TBTCl4～5w后,海水鳗鲡肝脏的谷胱甘肽-S-转移酶(GST)活性比对照组显著升高,而淡水鳗鲡组无明显变化;暴露于TBTCl或Phe3w后,淡水鳗鲡的血浆皮质醇水平显著升高,但海水鳗鲡组无明显变化.TBTCl和Phe对淡水或海水鳗鲡的血浆雌二醇(E2)水平均无明显影响.以上结果表明:有机锡和菲对鳗鲡的肝脏、脾脏有毒性作用,对皮质醇分泌具有内分泌干扰作用,且该效应与暴露时间和鱼体所处环境盐度有关.     

Orientation mechanisms in migrating European silver eel (Anguilla anguilla): Temperature and olfaction

The migration pattern of silver eel (Anguilla anguilla L.) in the Baltic is well documented from many tagging experiments. This particular investigation differed from previous taggings in that the background of all the eels used was the same. They came from a 1980 stocking programme and had been imported from France as glass eels. When migrating these stocked eels missed the outlet of the Baltic and continued in a south-westerly direction. They were still reported in this southern-most area more than 4 yr later. Indigenous eels turn northwards after passing the south of Sweden and leave the Baltic through the narrow Sound. Since the formerly stocked eels had never been in the Baltic, the absence of an imprinted olfaction cue may explain their orientation failure. Although lacking an imprinted olfaction cue eels were recaptured mainly in a very restricted area in the southwestern Baltic. This reflects another cue, genetic or imprinted during the Atlantic journey. This second cue, temperate, can also serve as the trigger which causes eels to stop migrating and to initiate spawning. The enclosed Baltic acts as a giant trap for eels from the huge stocking programmes undertaken there. As a consequence stocked eels probably contribute little to spawning stocks in the Sargasso Sea which may have contributed to the decrease in abundance of glass eels reported from western Europe in recent decades.     

Ontogenic change of gill chloride cells in leptocephalus and glass eel stages of the Japanese eel, <Emphasis Type="Italic">Anguilla japonica</Emphasis>

The development of gill chloride cells was examined in premetamorphic larvae (leptocephali) and juveniles (glass eels) of the Japanese eel, Anguilla japonica. Branchial chloride cells were detected by immunocytochemistry using an antiserum specific for Na⁺,K⁺-ATPase. The specificity and availability of the antiserum for the detection of Japanese eel chloride cells were confirmed by Western blot analysis. The chloride cells first appeared on the developing gill filaments in a mid larval stage of leptocephalus (32.2 mm). Both immunoreactivity and the number of chloride cells gradually increased as the fish grew to a late stage of leptocephalus over 54 mm. In glass eels just after metamorphosis, gill lamellae developed from the gill filaments, and a rich population of chloride cells was observed in the gill filaments. In glass eels collected at a coastal area, chloride cells were extensively distributed in the gill filaments. The chloride cell size decreased progressively in glass eels transferred from seawater (SW) to freshwater (FW), whereas there was no difference in cell number. In contrast, some Na⁺,K⁺-ATPase immunoreaction distinct from typical chloride cells was observed in the gill lamellae throughout FW-transferred fish, but disappeared in control fish maintained in SW for 14 days. These findings indicate that the gill and gill chloride cells developed slowly during the extremely long larval stage, followed by rapid differentiation during a short period of metamorphosis. The excellent euryhalinity of glass eels may be due to the presence of the filament chloride cells and lamellar Na⁺,K⁺-ATPase-immunoreaction, presumably being responsible for SW and FW adaptation, respectively.     

Occurrence of the semi-catadromous European eel Anguilla anguilla in the Baltic Sea

Strontium (Sr) and calcium (Ca) contents in the otoliths of yellow and silver European eels [Anguilla anguilla (L.)] collected from coastal waters of the Baltic Sea and a freshwater lake in Sweden were examined by wavelength dispersive X-ray spectrometry with an electron microprobe. The mean Sr/Ca ratios from elver check to otolith edge were significantly higher for the eels from coastal waters (5.39 ± 1.09‰) than for those from the lake (0.71 ± 0.89‰). The evidence indicates that European eels in the Baltic Sea do not necessarily migrate into freshwater streams during the growth phase. Received: 30 September 1999 / Accepted: 6 April 2000