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.     

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.     

Assessing patterns of hybridization between North Atlantic eels using diagnostic single-nucleotide polymorphisms

The two North Atlantic eel species, the European eel (Anguilla anguilla) and the American eel (Anguilla rostrata), spawn in partial sympatry in the Sargasso Sea, providing ample opportunity to interbreed. In this study, we used a RAD (Restriction site Associated DNA) sequencing approach to identify species-specific diagnostic single-nucleotide polymorphisms (SNPs) and design a low-density array that combined with screening of a diagnostic mitochondrial DNA marker. Eels from Iceland (N=159) and from the neighboring Faroe Islands (N=29) were genotyped, along with 94 larvae (49 European and 45 American eel) collected in the Sargasso Sea. Our SNP survey showed that the majority of Icelandic eels are pure European eels but there is also an important contribution of individuals of admixed ancestry (10.7%). Although most of the hybrids were identified as F1 hybrids from European eel female × American eel male crosses, backcrosses were also detected, including a first-generation backcross (F1 hybrid × pure European eel) and three individuals identified as second-generation backcrosses originating from American eel × F1 hybrid backcrosses interbreeding with pure European eels. In comparison, no hybrids were observed in the Faroe Islands, the closest bodies of land to Iceland. It is possible that hybrids show an intermediate migratory behaviour between the two parental species that ultimately brings hybrid larvae to the shores of Iceland, situated roughly halfway between the Sargasso Sea and Europe. Only two hybrids were observed among Sargasso Sea larvae, both backcrosses, but no F1 hybrids, that points to temporal variation in the occurrence of hybridization.     

A century of research on the larval distributions of the Atlantic eels: a re‐examination of the data

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 remain about the marine life histories of the Atlantic eels.     

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.     

Taxonomic revision,ecology and endangerment categorization of the Andean catfish Astroblepus ubidiai (Teleostei: Astroblepidae)

The European eel (Anguilla anguilla Linnaeus 1758) is a species typical for waters of Western Europe. Thanks to early expeditions on the Atlantic Ocean by the Danish biologist Johannes Schmidt who found small (<10mm) leptocephali larvae in the Sargasso Sea about 100 years ago, we have now a strong indication where the spawning site for this species is located. The American eel (Anguilla rostrata, LeSueur) also spawns in the Sargasso Sea. The spawning time and location of both species have been supported and refined in recent analyses of the available historical data. Subsequent ichthyoplankton surveys conducted by McCleave (USA) and Tesch (Germany) in the 1980s indicated an increase in the number of leptocephali <10 mm , confirming and refining the Sargasso Sea theory of Johannes Schmidt. Distinctions between the European and American eel are based on morphological characteristics (number of vertebrae) as well as molecular markers (allozymes, mitochondrial DNA and anonymous genomic-DNA. Although recognised as two distinct species, it remains unclear which mechanisms play a role in species separation during larval drift, and what orientation mechanism eels use during migration in the open sea. The current status of knowledge on these issues will be presented. The hypothesis that all European eel migrate to the Sargasso Sea for reproduction and comprise a single randomly mating population, the so called panmixia theory, was until recently broadly accepted. However, based on field observations, morphological parameters and molecular studies there are some indications that Schmidt’s claim of complete homogeneity of the European eel population and a unique spawning location may be an overstatement. Recent molecular work on European eel indicated a genetic mosaic consisting of several isolated groups, leading to a rejection of the panmixia theory. Nevertheless, the latest extensive genetic survey indicated that the geographical component of genetic structure lacked temporal stability, emphasising the need for temporal replication in the study of highly vagile marine species. Induced spawning of hormone treated eels in the aquarium was collective and simultaneous. In this work for the first time group spawning behaviour has ever been observed and recorded in eels. Studies in swim-tunnels indicate that eels can swim four to six times more efficiently than non-anguilliform fish such as trout. After a laboratory swim trial of eels over 5,500 km, the body composition did not change and fat, protein and carbohydrate were used in the same proportion. This study demonstrated for the first time that European eel are physiologically able of reaching the Sargasso Sea without feeding. Based on catches of newly hatched larvae, temperature preference tests and telemetry tracking of mature hormone treated animals, it can be hypothesised that spawning in the Sargasso Sea is collective and simultaneous, while presumably taking place in the upper 200 m of the ocean. Successful satellite tracking of longfin female eels in New Zealand has been performed to monitor migration pathways. Implementation of this new technology is possible in this species because it is three times larger than the European eel. In the future, miniaturisation of tagging technology may allow European eels to be tracked in time by satellite. The most interesting potential contribution of telemetry tracking of silver eels is additional knowledge about migration routes, rates, and depths. In combination with catches of larvae in the Sargasso Sea, it may elucidate the precise spawning locations of different eel species or groups. Only then, we will be able to define sustainable management issues by integrating this novel knowledge into spawners escapement and juvenile fishing quota.     

All roads lead to home: panmixia of European eel in the Sargasso Sea

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.     

On the geographic distribution and migration of I- and II-group eel larvae as studied during the 1979 Sargasso Sea Expedition

During the 1979 Sargasso Sea Expedition, 423 larvae ofAnguilla anguilla and 5 larvae ofA. rostrata were caught on three Atlantic transects and two cruises in the Sargasso Sea. Results of the identification of the larvae by myomere counts, and limits of the occurrence of I- and II-group larvae are presented. Four standard fishing depths are compared. A range shallower than 25 m was found to be the optimal fishing depth by night for both larval length groups. The geographic distribution of length group I was observed in central and eastern North Atlantic. Available data indicate a migration of these larvae in a north easterly direction. Length measurements of the II-group larvae taken from catches on the European continental slope during the same expedition support this assumption.     

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.     

Conclusive evidence for panmixia in the American eel

Eels are unique species in the biological world. The two North Atlantic eel species, the American eel (Anguilla rostrata) and the European eel (A. anguilla), occupy a broad range of habitats from the Caribbean to Greenland in the western Atlantic and from Morocco to Iceland in the eastern Atlantic, respectively. North Atlantic eels have a catadromous life cycle, spawning only in the Sargasso Sea and spending the majority of their lives in continental (fresh, brackish and coastal) waters. Despite such a wide distribution range, North Atlantic eels have been regarded as a textbook example of panmictic species. In contrast with the large amount of population genetic studies testing the panmixia hypothesis in the European eel, a relatively modest effort has been given to study the population structure of the American eel. In this issue of Molecular Ecology, Côté et al. ( 2013 ) present the most comprehensive American eel data set to date, which includes samples of different life stages obtained throughout all its distribution range in North America. Results show a total lack of genetic differentiation among samples and provide decisive evidence for panmixia in the American eel.     

Telemetric observations on the spawning migration of the eel (Anguilla anguilla) west of the European continental shelf

Synopsis In the Northern Bay of Biscay and west of the Iberian continental shelf five silver eels (Anguilla anguilla L.) have been tagged with ultrasonic transmitters and tracked 13 to 23 hours over a depth of 200 to 2500 m. Their mean direction from release to the final position of tracking was 288° and significantly closer to the direction of the Sargasso Sea (250° west) than silver eels tracked earlier in the North Sea (341°), possibly 260°. Four of the transmitters were equipped with pressure sensing devices capable of indicating depths of at least 400 m. Three eels tracked at night, during full moon, preferred mean depths of 125, 166 or 215 m. One eel chose a depth of 100 m during moonlight and 50 m after the setting of the moon. Major depth changes, usually occurring one per hour, ranged up to a maximum of 200 m at a maximum vertical speed of 0.6 m sec^–1; this is close to the eels' normal horizontal speed. At dawn all but one dived to a depth of 400 m or more. The eels generally swam below the thermocline and often crossed it.     

An update of the length–weight and length–age relationships of the European eel (Anguilla anguilla,Linnaeus 1758) in the Comacchio Lagoon,northeast Adriatic Sea,Italy

An update of length–weight and length–age relationships for yellow, silver and mixed population of European eels (Anguilla anguilla L.) of the Comacchio Lagoon, Italy, is provided in this study using data obtained in 2011. This historically important eel stock has undergone 99% feminization, probably due to the significant density reduction over the past three decades. The results show that these conditions did not affect the length–weight relationships, which approximate those of the late 1970s, but they do affect the length–age relationships, leading to faster maturation rates. These conditions lead to younger, longer and heavier silver eels before their migration to the Sargasso Sea.     

Comparison of electrophoretic and meristic characters of 0-group eel larvae from the Sargasso Sea

Marked differences between continental samples of American and European eels have been detected electrophoretically in allele frequencies at the MDH-2 locus. Starch gel electrophoresis carried out on board F. R. V. Anton Dohrn during the eel expedition to the Sargasso Sea in 1979 revealed a similar clear-cut genetic difference in a sample of 0-groupAnguilla leptocephali, thus confirming the classical theory of Schmidt (1932). The MDH-2 genotypes provide an additional diagnostic character for the distinction between youngA. anguilla andA. rostrata leptocephali. Species identification by biochemical genetic characters did not correspond with that by meristic characters (myomere numbers) in ca. 13 % of the specimens studied; this discrepancy mainly concerns leptocephali of theA. anguilla genotype. The results obtained are critically discussed.     

Are Lithuanian eels fat enough to reach the spawning grounds?

Stocks of the European eel Anguilla anguilla have been in a steep decline since the 1980s. Stocking of water bodies with juvenile eels captured in the wild to establish or enhance local populations has been a common practise in Europe for many decades. However, the degree of contribution by stocked eels to natural spawning capacity is poorly known and extensively debated. There have been suggestions that eels derived from stocking are less likely to contribute to the spawning stock due to a lack of navigational capability and lower fitness related to insufficiency of energetic resources. Results of the current study indicated that eels translocated long distances from the point of capture and released into inland waters in Lithuania are successfully undergoing the silvering process. A proportion of 23.7% (N = 27) among all migrating eels were described to be at the yellow (SI, SFII or SFIII) eel stage and downstream movements of these eels should be attributed to local movements, rather than spawning migration; 76.3% were assigned to the silver eel stage. This study suggests that 36.8% (N = 32) of downstream migrating silver eels of stocked origin had accumulated sufficient energetic resources for spawning migration and gonadal development and should be able to traverse the 7900-km distance to the presumptive spawning grounds in the Sargasso Sea. The rest of migrating silver eels (63.2%, N = 55) had insufficient energetic resources; the average potential swimming range of these eels was estimated to be 6135 ± 683 km.     

Progress report on the eel expedition of R.V. ‘Anton Dohrn’ and R.V. ‘Friedrich Heincke’ to the Sargasso Sea 1979

Synopsis This report presents preliminary results of the 1979 Sargasso Sea expedition from February to May 1979. Information is given on horizontal and vertical distribution of eel larvae and adults, adult eel tracking and pelagic trawling. Related matters such as electrophoretic studies on anguilliform larvae, feeding of eel larvae, predation on leptocephali, occurrence of other anguilliform larvae and hydrography are mentioned.     

The Distribution and Ecology of Ariosoma Balearicum (Congridae) Leptocephali in the Western North Atlantic

A total of 4589 leptocephali of the congrid eel, Ariosoma balearicum, were examined from 17 cruises in the western North Atlantic Ocean. Myomere counts made on 915 of these indicated there were two ranges of number of myomeres that appear to be associated with separate spawning populations. Those with the higher range (high count: 128–137) were consistently 70–100mm in length in the Sargasso Sea from February to April and 20–80mm in length in the northern Sargasso Sea and Gulf Stream from September to October. Those with the lower range (low count: 120–130) were rare in the northern and eastern Sargasso Sea where they had consistently greater lengths than high count leptocephali and were most abundant in the Florida Current and Providence Channel. The geographic distributions of size and myomere ranges in relation to hydrography provide strong support for the hypothesis that high count eels found along the South Atlantic Bight (SAB) migrate across the Florida Current to spawn in the northwest Sargasso Sea. This migratory pattern is similar to those of Anguilla rostrata and Conger oceanicus, which use the southern Sargasso Sea for development as larvae. However, the distribution of high count leptocephali suggests that they use the entire Sargasso Sea gyre as a development area as larvae before crossing the Florida Current and recruiting to the SAB. The low count eels inhabiting the Bahamas appear to spawn near the banks and their abundance in the Providence Channel and southwest Sargasso Sea suggests most are retained close to the Bahamas. These two distinct styles of spawning, distribution and recruitment of larvae are hypothesized to be related to the different hydrographic regimes of the two juvenile habitats and the resulting constraints on growth and recruitment of larvae. Vertebral and myomere counts reported from other areas suggest there are distinct populations in other regions of the North Atlantic Ocean.     

Larval Gnathostoma spinigerum Detected in Asian Swamp Eels,Monopterus albus,Purchased from a Local Market in Yangon,Myanmar

The present study was performed to determine the infection status of swamp eels with Gnathostoma sp. larvae in Myanmar. We purchased total 37 Asian swamp eels, Monopterus albus, from a local market in Yangon in June and December 2013 and 2014. All collected eels were transferred with ice to our laboratory and each of them was examined by the artificial digestion technique. A total of 401 larval gnathostomes (1-96 larvae/eel) were detected in 33 (89.2%) swamp eels. Most of the larvae (n=383; 95.5%) were found in the muscle. The remaining 18 larvae were detected in the viscera. The advanced third-stage larvae (AdL₃) were 2.3-4.4 mm long and 0.25-0.425 mm wide. The characteristic head bulb (0.093 × 0.221 mm in average size) with 4 rows of hooklets, muscular long esophagus (1.025 mm), and 2 pairs of cervical sacs (0.574 mm) were observed by light microscopy. The average number of hooklets in the 1st, 2nd, 3rd, and 4th rows was 41, 45, 48, and 51, respectively. As scanning electron microscopic findings, the characteristic 4-5 rows of hooklets on the head bulb, a cervical papilla, tegumental spines regularly arranged in the transverse striations, and an anus were well observed. Based on these morphological characters, they were identified as the AdL₃ of Gnathostoma spinigerum. By the present study, it has been confirmed for the first time that Asian swamp eels, M. albus, from Yangon, Myanmar are heavily infected with G. spinigerum larvae.     

Catadromous eels continue to be slippery research subjects

As adults, Atlantic eels (Anguilla rostrata in the Americas and Anguilla anguilla in Europe) are tubular slime-covered fish that spend most of their catadromous life-cycle in coastal environs before swimming far out to sea to reproduce, as part of an intergenerational migratory circuit that provides an interesting reversal of the pattern displayed by adult anadromous salmon that live mostly in the ocean but then migrate long distances to spawn in freshwater streams. Earlier genetic findings on Atlantic eels involved specimens collected across their broad continental ranges and generally indicated that conspecifics probably engage in panmictic or quasi-panmictic spawning,from which arise leaf-shaped leptocephaus larvae that then disperse back to coastal locations more or less at random with respect to the widespread geographical origins of the parental genes they carry. In this issue, Alset al. (2011) add exciting information about this peculiar life-history pattern of catadromous Atlantic eels by extending the genetic analyses to eel larvae collected from the Sargasso Sea, the oceanic area where both species spawn. Results help to confirm standard textbook wisdom that these catadromous eels are nearly unique in the biological world by having both broad geographical distributions and yet displaying intraspecific near panmixia.     

Egg size and the evolution of phenotypic plasticity in larvae of the echinoid genus Strongylocentrotus

Planktotrophic larvae grow by utilizing energy obtained from food gathered in the plankton. Morphological plasticity of feeding structures has been demonstrated in multiple phyla, in which food-limited larvae increase feeding structure size to increase feeding rates. However, before larvae can feed exogenously they depend largely on material contained within the egg to build larval structures and to fuel larval metabolism. Thus, the capacity for plasticity of feeding structures early in development may depend on egg size. Using the congeneric sea urchins Strongylocentrotus franciscanus and S. purpuratus, which differ in egg volume by 5-fold, I tested whether the degree of expression of feeding structure (larval arm length) plasticity is correlated with differences in the size of the egg. I experimentally manipulated egg size of S. franciscanus (the larger-egged species) by separating blastomeres at the 2-cell stage to produce half-sized larvae. I reared half-size and normal-size larvae under high and low food treatments for 20 days. I measured arm and body lengths at multiple ages during development and calculated the degree of plasticity expressed by larvae from all treatments. Control and unmanipulated S. franciscanus larvae (from ∼ 1.0 nl eggs) had significantly longer arms relative to body size and a significantly greater degree of plasticity than half-sized S. franciscanus larvae (from < 0.18 nl eggs), which in turn expressed a significantly greater degree of plasticity than S. purpuratus larvae (from ∼ 0.3 nl eggs). These results indicate that egg size affects larval arm length plasticity in the genus Strongylocentrotus; larger eggs produce more-plastic larvae both in an experimental and a comparative context. However, changes in egg size alone are not sufficient to account for evolved differences in the pattern of plasticity expressed by each species over time and may not be sufficient for the evolutionary transition from feeding to non-feeding.     

Diving activity of migrating silver eel with and without Anguillicola crassus infection

Infection with the swim bladder nematode Anguillicola crassus has been hypothesised to threaten the spawning migration success of the endangered European eel (Anguilla anguilla). To examine this assumption, we compared the swimming behaviour of one Anguillicola crassus infested eel in the North Sea and three parasite‐free eels in the Baltic using data recovered from data storage tags attached to migrating silver eels. In both areas, eel activity was characterized by frequent diving behaviour throughout the water column during the night, with reduced activity during the day. Despite substantial damage of the swim bladder, the behaviour of the infested eel from the North Sea was within the same range of migrating and diving activity parameters as the three parasite‐free eels from the Baltic Sea. All eels had a similar frequency distribution of descent or ascent speeds and a similar average horizontal migration speed. The diving speeds and dive ranges exclude the possibility that the eels were in continuous hydrostatic equilibrium during their migrations and suggests therefore that the role of the swim bladder in vertical migration is likely to be more complex than currently thought. Our results suggest that eels infested by Anguillicola crassus are capable of diving in a similar manner to uninfested eels during the first stretch of their spawning migration.