European eels (Anguilla anguilla) undertake a approximately 5000-kilometer (km) spawning migration from Europe to the Sargasso Sea. The larvae are transported back to European waters by the Gulf Stream and North Atlantic Drift. However, details of the spawning migration remain unknown because tracking eels in the Atlantic Ocean has, so far, eluded study. Recent advances in satellite tracking enable investigation of migratory behavior of large ocean-dwelling animals. However, sizes of available tags have precluded tracking smaller animals like European eels. Here, we present information about the swimming direction, depth, and migratory behavior of European eels during spawning migration, based on a miniaturized pop-up satellite archival transmitter. Although the tagging experiment fell short of revealing the full migration to the Sargasso Sea, the data covered the first 1300 km and provided unique insights.
European eels (Anguilla anguilla) spawn in the remote Sargasso Sea in partial sympatry with American eels (Anguilla rostrata), and juveniles are transported more than 5000 km back to the European and North African coasts. The two species have been regarded as classic textbook examples of panmixia, each comprising a single, randomly mating population. However, several recent studies based on continental samples have found subtle, but significant, genetic differentiation, interpreted as geographical or temporal heterogeneity between samples. Moreover, European and American eels can hybridize, but hybrids have been observed almost exclusively in Iceland, suggesting hybridization in a specific region of the Sargasso Sea and subsequent nonrandom dispersal of larvae. Here, we report the first molecular population genetics study based on analysis of 21 microsatellite loci in larvae of both Atlantic eel species sampled directly in the spawning area, supplemented by analysis of European glass eel samples. Despite a clear East-West gradient in the overlapping distribution of the two species in the Sargasso Sea, we only observed a single putative hybrid, providing evidence against the hypothesis of a wide marine hybrid zone. Analyses of genetic differentiation, isolation by distance, isolation by time and assignment tests provided strong evidence for panmixia in both the Sargasso Sea and across all continental samples of European eel after accounting for the presence of sibs among newly hatched larvae. European eel has declined catastrophically, and our findings call for management of the species as a single unit, necessitating coordinated international conservation efforts.
The spawning areas of the Atlantic freshwater eels were discovered about a century ago by the Danish scientist Johannes Schmidt who after years of searching found newly hatched larvae of the European eel, Anguilla anguilla, and the American eel, Anguilla rostrata, in the southern Sargasso Sea. The discovery showed that anguillid eels migrate thousands of kilometers to offshore spawning areas for reproduction, and that their larvae, called leptocephali, are transported equally long distances by ocean currents to their continental recruitment areas. The spawning sites were found to be related to oceanographic conditions several decades later by German and American surveys from 1979 to 1989 and by a Danish survey in 2007 and a German survey in 2011. All these later surveys showed that spawning occurred within a restricted latitudinal range, between temperature fronts within the Subtropical Convergence Zone of the Sargasso Sea. New data and re-examinations of Schmidt's data confirmed his original conclusions about the two species having some overlap in spawning areas. Although there have been additional collections of leptocephali in various parts of the North Atlantic, and both otolith research and transport modelling studies have subsequently been carried out, there is still a range of unresolved questions about the routes of larval transport and durations of migration. This paper reviews the history and basic findings of surveys for anguillid leptocephali in the North Atlantic and analyses a new comprehensive database that includes 22612 A. anguilla and 9634 A. rostrata leptocephali, which provides a detailed view of the spatial and temporal distributions and size of the larvae across the Atlantic basin and in the Mediterranean Sea. The differences in distributions, maximum sizes, and growth rates of the two species of larvae are likely linked to the contrasting migration distances to their recruitment areas on each side of the basin. Anguilla rostrata leptocephali originate from a more western spawning area, grow faster, and metamorphose at smaller sizes of <70 mm than the larvae of A. anguilla, which mostly are spawned further east and can reach sizes of almost 90 mm. The larvae of A. rostrata spread west and northwest from the spawning area as they grow larger, with some being present in the western Caribbean and eastern Gulf of Mexico. Larvae of A. anguilla appear to be able to reach Europe by entering the Gulf Stream system or by being entrained into frontal countercurrents that transport them directly northeastward. The larval duration of A. anguilla is suggested to be quite variable, but gaps in sampling effort prevent firm conclusions. Although knowledge about larval behaviour is lacking, some influences of directional swimming are implicated by the temporal distributions of the largest larvae. Ocean-atmosphere changes have been hypothesized to affect the survival of the larvae and cause reduced recruitment, so even after about a century following the discovery of their spawning areas, mysteries still rema...
The two primary ways that species respond to heterogeneous environments is through local adaptation and phenotypic plasticity. The American eel (Anguilla rostrata) presents a paradox; despite inhabiting drastically different environments [1], the species is panmictic [2, 3]. Spawning takes place only in the southern Sargasso Sea in the Atlantic Ocean [1]. Then, the planktonic larvae (leptocephali) disperse to rearing locations from Cuba to Greenland, and juveniles colonize either freshwater or brackish/saltwater habitats, where they spend 3-25 years before returning to the Sargasso Sea to spawn as a panmictic species. Depending on rearing habitat, individuals exhibit drastically different ecotypes [4-6]. In particular, individuals rearing in freshwater tend to grow slowly and mature older and are more likely to be female in comparison to individuals that rear in brackish/saltwater [4, 6]. The hypothesis that phenotypic plasticity alone can account for all of the differences was not supported by three independent controlled experiments [7-10]. Here, we present a genome-wide association study that demonstrates a polygenic basis that discriminates these habitat-specific ecotypes belonging to the same panmictic population. We found that 331 co-varying loci out of 42,424 initially considered were associated with the divergent ecotypes, allowing a reclassification of 89.6%. These 331 SNPs are associated with 101 genes that represent vascular and morphological development, calcium ion regulation, growth and transcription factors, and olfactory receptors. Our results are consistent with divergent natural selection of phenotypes and/or genotype-dependent habitat choice by individuals that results in these genetic differences between habitats, occurring every generation anew in this panmictic species.
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.
We critically review four potential causes of a drastic decline in juvenile American eels, Anguilla rostrata, recruiting to Lake Ontario (81-fold decline from 1985 to 1992) and in juvenile eel densities in tributaries to the Gulf of St. Lawrence. Silver eels from the St. Lawrence River were much more contaminated and had a high prevalence of deformities and lesions than a reference stock although they were on average less contaminated in 1990 than in 1982 (1.12 versus 4.54 μgg−1 for PCB; 0.025 versus 0.07 μg∙g−1 for mirex). Lethal toxicity from chemical contamination has been known to occur in St. Lawrence River eels for the past 25–30 yr. Major habitat modifications in the St. Lawrence took place in the 1950's (St. Lawrence Seaway and hydroelectric dams), about 30 yr before recruitment started declining; this long delay argues against these perturbations being primary causes of the decline. There is little evidence that commercial fishing and oceanographic changes caused the decline. Overall, we conclude that we do not know what caused the pronounced recruitment decline. We propose research avenues and hypotheses that may advance understanding and emphasize that because of panmixia, the recruitment decline could be species wide.
1991. Usefulness of Fourier analysis of otolith shape for Atlantic mackerel (Seornbea scombsus) stock discrimination. Can. ). Fish. Aquat. Sci. 48: 296-302.We compared shapes of Atlantic mackerel (Seornber scombrus) sagittae between the two contingents (i .e. spawning groups) from the Northwest Atlantic and between the stocks from the Northwest Atlantic and the North Sea to evaluate whether otolith shape could differentiate between the two contingents in a mixed fishery. We quantified shapes with the Fourier series, an objective and rapid method which decomposes a shape's outline into a series of sinusoids. To detern-sine a correct way to compare contingentslstocks, we first assessed four intracontingent effects on otolith shapes. Age and year-class effects were significant, while sex and bilateral position effects were not. This temporal instability in shapes indicates that confounding effects of age and year-class on otolith shapes need to be assessed carefully before drawing conclusions on stocl< structure. It also shows that comparative studies of otolith shapes with Fourier descriptors are not useful for mackerel contingent discrimination. The power of discriminant functions to correctly classify test mackerel samples separated by age and sampling year, on the basis of otolith shape, was better for comparisons between the stocks than for those between the contingents. Nous avons compare la forme de sagittae de maquereau bleu (Scsmbes scombaus) entre Bes deux contingents (i.e. group-reproducteurs) du nord-ouest de I'Atlantique et entre les stocks dea nord-ouest de I'Atlantique et de la mer du Nord afin de determiner s'il est possible de diffprencier les deux contingents sur la base des formes d'otolithes dans une peche rniwte. Nous avons quantifik les formes A l'aide des sgries de Fourier, une rnkthode objective et rapide qui d6csrnpose le contour d'une forme en une skrie de sinussides. Afin de dkterrniner une f a~o n correcte de cornparer Ies csntingents/stscks, nous avons d'absrd 6vaIu6 quatre effets intra-contingents sur tes formes d'otolithes. Les effets d'sge et de classe d'sge sur les formes d'stolithes etaient significatifs alors que les effets de sexe et de position bilatkrale ne It6taient pas. Cette instabilitk ternporelle des formes indique que les effets d'age et de classe d'2ge doivent &re soigneusement 6valu6s avant de tirer des conclusions sear la structure de stock et d6montre que des comparaisons de formes d'otolithes avec les descripteurs de Fourier ne sont pas utiies pour la discrimination des contingents de maquereau. La puissance de fonctions discriminantes 2 classifier correctement des kchanti l Ions tests de maquereau separks par 2ge et ann6e d'kchanti llonnage sur la base des forrnes d'otslithes etait meilleure pour les comparaisons entre les stocks que pour celles entre les contingents.
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