Using otolith microchemical analysis to investigate the importance of brackish bays for pike (Esox lucius Linnaeus, 1758) reproduction in the southern Baltic Sea

The brackish Baltic Sea is inhabited by several primary freshwater fish species. The pike (Esox lucius) is the largest of these species and acts as the top predator in the marine food web of the Baltic Sea. Prior studies have shown the importance of brackish bays as spawning grounds for the Baltic Sea pike. To identify the spawning habitat of the pike population around the island of Rügen (southern Baltic Sea)—either freshwater streams flowing into the Baltic or brackish lagoons of the Baltic—we used the ICPMS method to quantify the strontium to calcium (Sr:Ca) ratios in sagittal otoliths of 79 adult pike. In total, only six per cent (n = 5) of the analysed Baltic Sea individuals were classified as being of freshwater origin. Ninety‐four per cent (n = 74) of analysed pike were classified as having hatched in brackish water. Compared to previous studies on Baltic Sea pike, our sample shows a high proportion of pike that is of brackish origin: this highlights the adaptive capacity of this species with regard to its ability to reproduce in different salinities. Our data highlight the importance of brackish bays for pike reproduction in the southern Baltic Sea, even if pike live on the upper limit of their salinity tolerance in this area. This is an important finding considering decreases in commercial catches in the last 30 years indicating a population decline supposedly related to recruitment failures.     

Genetic population structure of pike (Esox lucius Linnaeus, 1758) in the brackish lagoons of the southern Baltic Sea

The Baltic Sea is a unique ecosystem and the largest body of brackish water in the world. It is inhabited by several originally freshwater fish species, one of them being pike (Esox lucius), which is adapted to the mesohaline conditions of the southern Baltic Sea and colonises its south‐western bays and lagoons. Using twelve microsatellite markers, we analysed the genetic structure of 407 Baltic Sea pike collected from ten locations in the coastal zone of the southern Baltic region and adjacent rivers and nearby lakes. We used traditional F‐statistics and a Bayesian approach implemented in STRUCTURE to investigate spatial structuring. Our results show an overall isolation by distance pattern with two distinct coastal clusters. The border area between the two clusters is characterised by a sharp salinity leap, indicating that the dispersal barrier is not physical, but physiological in nature. A third cluster was formed by three lake populations, presumably as a result of past stocking measures. Additional otolith microchemical studies demonstrate the brackish origin and residency of the majority of the pike from the coastal lagoons, supporting previous studies which suggest that the anadromous pike “ecotype” is scarce in this area of the Baltic Sea. Our results contribute to current knowledge on the population genetics of Baltic Sea pike in a previously unsampled area.     

Do Eurasian minnows (Phoxinus phoxinus L.) inhabiting brackish water enter fresh water to reproduce: Evidence from a study on otolith microchemistry

Dynamics of coastal fish assemblages in the Baltic Sea are still rather poorly understood. In particular, little information is available on migrations and movements of the small‐bodied littoral fish species like Eurasian minnow (Phoxinus phoxinus L.). Minnow is considered typically as a river and lake species, but it also inhabits brackish coastal waters of the Baltic Sea. In this study, we investigated movement patterns of brackish water inhabiting minnows using otolith microchemistry. Fish were collected from four different sites (from two bays and two stream mouths) around Saaremaa Island. The results indicated that at least three distinct migration patterns exist: (i) seawater residency, (ii) fast springtime migrations to fresh water or to bays with lower salinities (areas near stream mouths) and (iii) prolonged migrations to fresh water (some cases overwintering in streams). Migration patterns listed above were not evenly distributed among sites, and some individuals did not migrate to fresh water in every year. None of the analysed fish were freshwater residents. Additionally, potentially ontogenetic effects on Mn and Sr concentrations were observed in the otolith core regions, which may have important implications for the interpretation of otolith chemistry data. The overall findings of this study demonstrate that much more complex migration patterns exists in brackish water inhabiting minnow populations than previously thought and at least some individuals do enter freshwater to reproduce.     

High salinity tolerance in eggs and fry of a brackish Esox lucius population

Abstract Knowledge on the biology and physiology of pike, Esox lucius L., populations inhabiting saline environments is scarce. An experimental setup was used to examine egg development and fry behaviour and growth under varying salinity levels in a brackish‐water pike population from the western Baltic Sea. Eggs and fry developed at 8.5 psu, which is higher than hitherto reported for other populations. Fry exhibited stress behaviour and reduced growth when subjected to salinities above 13 psu. This indicates that early life stages of E. lucius tolerate ambient salinity conditions equivalent to the natural environment in its nursery area in the Stege Nor lagoon. The population should therefore not be dependent on access to freshwater environments during spawning. However, the results also suggest that the population’s natural environment is on the margin of its physiological capacity. The results have significance for management of brackish‐water E. lucius populations, e.g. in relation to stocking strategies.     

Elemental fingerprinting in otoliths reveals natal homing of anadromous Baltic Sea pike (Esox lucius L.)

We examined the element pattern in the otoliths of a migratory fish species that inhabit the coastal areas in the brackish of the Baltic Sea. The northern pike (Esox lucius) show migratory behaviour, spawning in streams and rivers and foraging in the sea. We examined spawning migration in four nearby streams in the south‐west part of the Baltic. Otolith analysis by microPIXE revealed unique elemental patterns (Sr, Zn, Br, Co and Mn) for the juveniles in each of the different streams. The strontium signal in the otolith of the juveniles was used as an indicator of freshwater origin and the time spent in the stream. Adult pike in their migrating spawning phase were caught in each of the streams. The elemental composition in otoliths in their freshwater phase (using juvenile pike in the streams as references) was determined. A principal component analysis showed that the elemental fingerprint during the freshwater phase several years back in time was similar for the adult fish and for juveniles inhabiting the stream today. The results indicated natal homing of the adults to a specific stream, a conclusion that was strengthened by the fact that marked fish returned to spawn over consecutive years. Anadromous pike in the Baltic Sea may thus be divided in subpopulations. The results of the study may have implications for fishery management, as pike in the Baltic Sea cannot be seen as homogenous population.     

Genetic differentiation between and within ecotypes of pike (Esox lucius) in the Baltic Sea

1. Aquatic systems often lack physical boundaries for gene flow, but ecological and behavioural barriers can form surprisingly fine spatial scale genetic patterns that challenge traditional, large scale management. To detect fine spatial scale structures, understand sources of intraspecific diversity, and design appropriate management plans requires identification of reproductively isolated units.
2. This study reports on genetic differentiation in pike (Esox lucius) within a coastal area stretching 55 km from south to north in the central Baltic Sea. Pike is here an economically and ecologically important top predator that has declined in abundance. However, population structures have mostly been studied on large spatial scales, and without considering the potential for genetic divergence between the sympatric anadromous fresh water and the resident brackish water spawning ecotypes.
3. To this end, 487 individuals from the east coast of Sweden and the island of Öland, representing sympatric anadromous and resident spawning individuals, categorized to ecotype based on spawning location or otolith microchemistry, were genotyped for 10 microsatellites and used to test for divergence between ecotypes. Furthermore, divergence between regions (island/mainland), neighbouring spawning locations (n = 13) and isolation by distance within and between regions were evaluated for the anadromous ecotype.
4. The results revealed strong genetic differences between regions, between spawning locations separated by as little as 5 km and the first evidence of genetic differentiation between resident and anadromous ecotypes; despite a high dispersal capacity of pike and a high connectivity within the study area. The signatures of isolation by distance indicated that connectivity among populations differed between regions, probably reflecting availability of spawning habitats.
5. To safeguard against the challenges and uncertainties associated with environmental change, adaptive conservation management should aim to promote high intra‐population functional genetic diversity without compromising the continued integrity and coexistence of the different ecotypes and of locally adapted sub‐populations.

     

Abiotic triggers for sea and river lamprey spawning migration and juvenile outmigration in the River Rhine,Germany

The activity patterns of migrating lampreys (sea lamprey Petromyzon marinus and river lamprey Lampetra fluviatilis) are poorly understood. This study uses extensive screening data from the cooling water intake of a nuclear power plant on the River Rhine, Germany, gathered on 1308 days over 46 consecutive months between 2011 and 2014. More than 34,000 lampreys were retrieved, of which the great majority (99.9%) were outmigrating nonfeeding postmetamorphic individuals. The main factors controlling outmigration were season, water temperature and river discharge. The bulk of downstream migration was observed each year in spring at water temperatures between 9 and 12°C, with earlier migrations during significant discharge peaks between December and February. The migration activity of postmetamorphic individuals declined during summer and autumn, regardless of discharge. Turbidity increased the occurrence of postmetamorphic lampreys and supported diurnal migration, which accounted for nearly 20% of all lamprey records. In winter, outmigrating individuals of both species exhibited significantly longer body lengths than those detected during other seasons. Haematophagus postmetamorphic sea lampreys were also retrieved (n = 28) and represent the most upstream records of parasitically feeding sea lamprey detected thus far. Adult lampreys migrating upstream were found mainly in spring, but in very low numbers, indicating high prespawning mortalities. The study points to a challenging future for the protection of lampreys in this area: the long duration and the round‐the‐clock nature of the observed migration demand flexible solutions to realise safe operational windows for water intake stations and to minimise impacts on juvenile lampreys.     

Vertical movements of coastal pike (Esox lucius)—On the role of sun basking

Knowledge of patterns and drivers of the spatiotemporal distribution of top predatory fish is key to understand ecological dynamics and to successful management. Here, we integrated field and laboratory approaches to study vertical movements of pike (Esox lucius) in relation to season, light regimes and body temperature. We tagged pike from the Baltic Sea with data storage tags during spawning migration and retrieved them during migration the following years to obtain high-resolution data from full year of movements. The results showed seasonal and diel patterns of activity and body temperature that conformed to distinct patterns of crepuscular activity and diel vertical migrations. The latter manifested as two different patterns, either a stationary phase in the surface water during day followed by night-time in deeper water or vice versa. The occurrence of these two behaviours varied among individuals and within individuals among seasons. Diel vertical migration has previously not been described for this shallow-dwelling species, but was a common and consistent behaviour among individuals in this study. We suggest that the function of the daytime surface behaviour in pike is to increase body temperature through sun basking. This thermoregulatory role of surfacing was supported by the laboratory study where individuals sought the surface layer, exposed themselves to infrared light and thereby attained body temperatures in excess of ambient water. These results support sun basking as a mechanism for heat gain and further suggest that access to sunlight in the surface layer could be an important driver of vertical migrations.     

Meta‐analysis of growth rates for a circumpolar fish,the northern pike ( Esox lucius), with emphasis on effects of continent,climate and latitude

For circumpolar species, little is known on how somatic growth rates can vary at large, transcontinental spatial scales. In this study, a meta‐analysis of growth rates was conducted for northern pike ( Esox lucius) across North America and Eurasia. Growth rates of northern pike did not differ between North American and ‘coastal Eurasian’ pike (e.g., UK, Ireland, Sweden), while growth rates for both of these groups were significantly higher compared to ‘inland Eurasian’ pike (mainly in Russia). There was no difference in growth between lentic and lotic habitats on either continent. In North America, pike growth was positively correlated with temperature, but in Eurasia, pike growth correlated poorly with most climatic variables. Similarly, maximum longevity in pike populations was significantly predicted by latitude in North America, but not in Eurasia. After standardising annual pike growth by the thermal opportunity for growth, a highly significant countergradient growth relationship was found for North American pike, while a significant, but considerably less predictive countergradient growth relationship was found for Eurasian pike. This study provides novel insights into the ecology of a circumpolar species and how populations function at extraordinarily large spatial scales. First, pike appear to be cosmopolitan across hydrologic habitats having fast or slow growth in either lentic or lotic environments. Secondly, continental‐scale differences in pike growth rates are suggestive of major genetic and life‐history differences. Finally, variable climate–growth relationships and countergradient growth patterns indicate that global climate change is likely to affect circumpolar fishes like pike in complex, nonlinear ways.     

Effects of salinity on growth and mortality of migratory and resident forms of Eurasian perch in the Baltic Sea

Abstract – In the coastal areas of the Baltic Sea, there are two forms of perch (Perca fluviatilis). One of the forms is migratory and spawns in streams entering the Baltic Sea. The other form is resident and spawns in brackish water. Both forms utilise the coastal habitat for foraging. We examined the spawning success of the two forms in fresh and brackish water (7‰, equal to salinity in the south Baltic Sea). The experiments showed that hatching success was equally high in freshwater and in brackish water despite female origin. The survival of yellow‐sac and free swimming fry was significantly reduced in brackish water, which was independent if the fish was of migratory or brackish resident origin. Further, growth rate of perch fry was severely reduced in brackish water. The results indicate that perch has not developed any tolerance to brackish water in the young life stages. The migratory life strategy of perch can thus be explained by higher survival of fry in freshwater.     

Behaviour,escapement and mortality of female European silver eels within a regulated lowland river draining into the Baltic Sea

Migration patterns of 146 female silver eels on their way through fresh and brackish water stretches of the Warnow River into the Baltic Sea were studied using acoustic telemetry. The majority of eels (77%) regained their expected downstream migration behaviour after a median rehabilitation time of 2 days. Positive rheotactic behaviour was observed for 23% of tagged eels after release, probably influenced by experimental procedures. The observed silver eel escapement rate was 63%. Two migratory groups were classified by significantly different migratory speed, time of residence in brackish waters and biometric silvering characteristics. Although the classification of silvering stages is based on these measures, the proportions of these stages did not reflect the observed differences in migratory behaviour. In the context of a permanent silver eel descent in the freshwater part of the Warnow River, a permanent escapement was likewise occurring into the Baltic Sea.     

Hatching time effect on the intra‐spawning larval morphology and growth in Northern pike (Esox luciusL.)

Based on the analysis of 11 morphometric variables of body (total length, body area and perimeter, myotome height and eye diameter) and yolk sac (length, height, area, perimeter and volume) of pike larvae, the aim of this study was to evaluate how larval size at hatching and growth of larvae hatched from single egg batches vary according to three hatching times: early, mid and late. Hatching time structures strongly pike larval morphometrics. Early hatched larvae have smaller body sizes at hatch, faster growth and higher yolk use efficiency than late hatched ones. Early hatched larvae seem to be premature and hatch at precocious developmental stage whereas late hatched individuals continue their growth within the egg shell and hatch at larger size but with lower reserves (yolk). A compensatory growth phase was observed for the early hatching pike larvae particularly during the first 5 days post hatch. Consequently, no significant difference in body parameters was recorded from day 10 post hatching whatever the hatching time. The higher growth accomplished by early hatched larvae may be related to a particular metabolic activity that converts more efficiently yolk into body tissues.     

Comparison of growth and reproduction performance of broodstock Pacific white shrimp Litopenaeus vannamei reared in oceanic and brackish water

This study provides a preliminary evaluation of feasibility of brackish water breeding. Shrimps were initially cultured at brackish water. Shrimps were reared at different stages (5, 6, 7 and 8 months, respectively), and 300 female shrimps were selected from each stage for growth at an enhanced salinity of 30 ppt. When the shrimps were sexually mature, the reproductive characteristics (mating rate, egg number and nauplii number) of the broodstock shrimps, quality of nauplii (malformation rate of nauplii) and quality of postlarvae (survival rate and desalination stress) were tested to evaluate of the effects of brackish water (10 ppt) on these shrimps. Oceanic water (30 ppt) for rearing shrimps was used as control group. Histopathological analysis of gonads and hepatopancreases from oceanic and brackish water shrimp was performed. The results showed that shrimps cultured for a maximum of 8 months in brackish water remained sexually mature. There were no significant differences regarding the quality of shrimps in the oceanic water and brackish water. However, the brackish water cultured shrimps were easily adapted to salinity changes. Our results demonstrate the good feasibility of the present scheme, in which the broodstock shrimps were cultured at a salinity of 10 ppt.     

Site fidelity,diel and seasonal activities of yellow‐phase Japanese eels (Anguilla japonica) in a freshwater habitat as inferred from acoustic telemetry

Japanese eels (Anguilla japonica) perform large‐scale oceanic migrations between their spawning ground and growth habitats in continental waters during life history. However, between these migrations, they spend most of time in growth habitats such as rivers. To investigate the diel and seasonal activities, homing behaviour and home range of yellow‐phase Japanese eels in the lower reach of the Tone River, we tracked them throughout a year using a fine‐scale positioning system (VPS) based on acoustic telemetry. The tracked eels were generally nocturnal, but not exclusively. They were mainly mobile from spring to autumn, with little or no activity observed during winter. A transport‐release experiment showed that most eels returned to their original capture area within 13 days after release. The eels had very small home ranges (mean ± SD = 0.085 ± 0.068 km²), core areas (0.014 ± 0.014 km²) and linear home ranges (744 ± 268 m). They also tended to be distributed on one particular side of the river (right or left bank) and in one particular shore type (revetment or vegetation), rarely moving from one to the other. This study provides evidence for nocturnal, dormancy, homing behaviours, limited habitat use and small home range size in Japanese eels. The eels clearly showed strong fidelity to a “familiar” site, which contrasts with the long distances travelled during upstream and downstream migration phases in the river, and during spawning migrations in the ocean.     

Movement patterns of adult pike (Esox lucius L.) in a Belgian lowland river

Northern pike, Esox lucius, needs different habitats to survive and reproduce and thus depends on the availability and accessibility of these habitats. To efficiently manage pike, information is needed on its spatial and temporal patterns of migration. In this study, we investigated the occurrence of adult pike migration and which environmental variables influenced migration. From December 2010, we followed 15 pike for 1 year by use of radio telemetry in the River Yser, a typical lowland river characterised by anthropogenic impacts such as artificial embankments. Pike migrated most in February and March, which could indicate they frequented spawning habitat in this period. Four environmental variables significantly affected pike migration, ranging from the location where pike were observed (strongest effect), over water temperature and flow to diel water temperature change (weakest effect). The relation between migration and the location where pike were observed could demonstrate that pike preferred specific regions in the river. Increasing water temperature triggered migration for both sexes, and males started migrating at lower temperatures than females, which suggests that males start migrating earlier. This was the only substantial difference observed between male and female pike migration. The results suggest that migration was inhibited by high flow, as no migration was observed at high flow. River managers can use this information to efficiently manage their pike populations, for example, by removing or temporarily opening hydraulic structures like valves, weirs and sluices. This may facilitate access to suitable habitats at moments pike needs these habitats to fulfil its life cycle.     

Migration of eels tagged in the Baltic Sea and Lake Mälaren—in the context of the stocking question

Eels (Anguilla spp.) are in decline worldwide, and the signs of reduced recruitment have been observed in continental Europe since the early 1970s. To aid recovery of the European eel, stocking is used by many European countries as a management option. In this study, tagging experiments were conducted to follow eel migration from Lake Mälaren and four sites along the Swedish east coast in the Baltic Sea. The recaptured tagged eels were retrieved from fishermen, allowing for the opportunity to investigate their origin (brackish water, stocked in freshwater or a mix in between) by otolith microchemistry and to assess for morphological differences after tagging. Several changes took place; for example, eye index increased while weight and condition decreased with migrated distance and time until recapture. In Lake Mälaren, the majority of tagged eels did not migrate out of the outlets, irrespective of their origin. Most of them were caught in the opposite direction and continued to be caught in the lake 1–3 years after tagging, with significant weight losses. Overall, overwintering is suggested to be an inferior option, but it is uncertain whether this is a natural behaviour or a result of translocation and restocking. For coastal eels, origin had no effect on migratory behaviour; a majority of the tagged eels migrated towards the outlet of the Baltic Sea. Interestingly, a minority of the recaptured eels originated from stocked fish. Instead, recaptures were dominated by natural immigrants that had spent most of their lives in brackish waters.     

An additional life-history tactic of masu salmon: Migration of parr to coastal habitats

Alternative life-history tactics of masu salmon Oncorhynchus masou are well documented. Subsequent to the freshwater parr stage (age ≥ 1 + years), an anadromous form migrates to the sea after smolting, while a resident form matures without seaward migration. In addition to this typical migratory dimorphism, anecdotal reports based on field observations have indicated that some underyearling masu salmon use estuarine waters. However, no empirical evidence indicates saltwater utilisation and subsequent survival in the early parr stage. Here, we used otolith microchemistry to examine whether a portion of masu salmon parr in northern Japan enters coastal habitats. The otolith Sr:Ca ratios of most juveniles collected from six rivers had consistently low values, indicating that masu salmon parr inhabiting these rivers stay only in freshwater. In contrast, in individuals from a steep-gradient river the Sr:Ca ratios increased at about a 200-µm distance from the otolith core. These results suggest that some masu salmon parr might use brackish water or sea water temporarily. In addition, three masu salmon parr were found in another steep river where a culvert located only ten metres from the river mouth completely blocked upstream migration for spawning. The Sr:Ca ratios in these fish increased at about >200 µm from the otolith core, indicating the parr had immigrated to the non-natal river from the sea. Such flexible behaviour at an early life stage may contribute to the spatial expansion of masu salmon, and the movement could moreover help to stabilise its population dynamics.     

Newcomers in the Baltic Sea: an attempt to trace the origins and whereabouts of thicklip grey mullet <Emphasis Type="Italic">Chelon labrosus</Emphasis>

In recent years, thicklip grey mullet Chelon labrosus has shown increasing expansion of its native habitats in the north-eastern Atlantic into northerly adjacent areas including the North Sea and the brackish Baltic Sea. Despite the regular annual and seasonal occurrence of C. labrosus in the western Baltic during the warm months, nothing is known of the origin or whereabouts of the mullet during the cold season. As different possible migration scenarios can be considered, we performed otolith microchemistry analyses on specimens from the western Baltic Sea to identify the origin of this nonindigenous species. Comparison with North Sea samples revealed common habitat preferences and underlined the highly euryhaline nature of C. labrosus in different recently occupied habitats. Occasional fluctuations of Sr/Ca ratio along the growth axis suggest periodical migration between waters of different salinities but did not reveal distinct migration pathways.     

What goes up must come down: Diel vertical migration in the deep‐water sablefish (Anoplopoma fimbria) revealed by pop‐up satellite archival tags

The sablefish (Anoplopoma fimbria) is a long‐lived species with wide distribution throughout the North Pacific Ocean. While adult sablefish are considered a deep‐water fish, diet analyses suggest that they undergo vertical migrations that could be related to prey movement and feeding. Pop‐up satellite archival tags (PSATs) were used to observe the fine‐scale depth selection behavior of adult sablefish tagged off the Washington coast during the summer from June to August. Tags were physically retrieved after they surfaced using direction‐finding equipment so that complete datasets over the entire deployment were obtained from 14 tags. PSATs that recorded depth and temperature every 4 min during the deployment confirm that sablefish inhabit depths of 750 m or greater. However, a majority of the tagged fish underwent extensive vertical migrations that averaged 254.4 m overall and occurred at a 24 hr periodicity. Variations were observed among individuals in the amount of the deployment during which vertical migrations occurred, ranging from 12.37% to 63.48% of the time. During the vertical migration, fish ascended towards the surface at night and descended prior to daylight (i.e., diel vertical migration). Sablefish generally inhabited temperatures of 5°C but during the vertical migrations were found at temperatures from 6 to 10°C. Sablefish are opportunistic feeders with a large proportion of their diet being fish, euphausiids and cephalopods. Because these prey items also exhibit diel vertical migrations, it is possible that the vertical migratory behavior displayed by the sablefish was in response to the movements or the location of their prey.     

Dispersal strategies of juvenile pike (Esox lucius L.): Influences and consequences for body size,somatic growth and trophic position

Individual variability in dispersal strategies, where some individuals disperse and others remain resident, is a common phenomenon across many species. Despite its important ecological consequences, the mechanisms and individual consequences of dispersal remain poorly understood. Here, riverine Northern pike (Esox lucius) juveniles (age 0+ (young-of-the-year) and 1+ years) were used to investigate the influence of body size and trophic position (at capture) on the dispersal from off-channel natal habitats and the subsequent consequences for body sizes, specific growth rate and trophic position (at recapture). Individuals that dispersed into the river (“dispersers”) were not significantly different in body size than those remaining on nursery grounds (“stayers”). For trophic position, 0+ dispersers were of significantly lower trophic position than stayers, but with this not apparent in the 1+ fish. Following dispersal into the river, the dispersers grew significantly faster than stayers and, on recapture, were significantly larger, but with no significant differences in their final trophic positions. Early dispersal into the river was, therefore, not associated with early dietary shifts to piscivory and the attainment of larger body sizes of individuals whilst in their natal habitats, contrary to prediction. These results suggest that despite an increasing risk of mortality for individuals dispersing early from natal areas, there are long-term benefits via elevated growth rates and, potentially, higher fitness. Such early dispersal behaviour could be driven by early competitive displacement.