Ontogenetic changes in habitat use by whitefish,Coregonus lavaretus

Synopsis The whitefish, Coregonus lavaretus, in the lake Mj?sa exhibited two niche changes during their life cycle. Juveniles (< 25 cm body length) were confined to the shallow (0–30 m) epibenthic zone. Medium sized whitefish (25–35 cm body length) expanded their habitat use to include the deep (30–90 m) epibenthic zone as well as the pelagic zone. From a body length of 35 cm, habitat use was restricted to the deep epibenthic zone. Small fish in the shallow epibenthic zone ate small and medium-sized prey (zooplankton, insect larvae and surface insects). Medium-sized fish in this zone were in addition feeding on the larger amphipod, Pallasea quadrispinosa. In the pelagic zone, the diet of medium-sized whitefish was dominated by zooplankton, although some larger prey like surface insects and age-0 smelt, Osmerus eperlanus, were also eaten. In the deep epibenthic zone, the diet of both medium-sized and large (< 35 cm) whitefish consisted mainly of the large prey P. quadrispinosa.     

Downstream displacement and life-history traits of whitefish,Coregonus lavaretus,in a Norwegian river

Synopsis Each year from 1976 to 1985 a part of the freshwater resident whitefish,Coregonus lavaretus, in Liavatn, south-western Norway, was displaced downstreams into the sea. In the sea, they seemed to die within a short period of time. The fish appeared to be flushed downstream by autumn freshets. In age and size, but not sex, the descenders seem representative of the fish in the lake. Females were the predominant sex in the river, whereas the sex ratio did not differ from unity in the lake. Young fish were fast-growing, but the growth rate levelled off at an age of 2 years and an asymptotic length of ∼30 cm. Males matured mainly at age 1+ and females at age 2+. The gonadosomatic index decreased with age, whereas the amount of energy allocated to gonadal tissue relative to somatic tissue increased with age     

Fecundity of whitefish (Coregonus lavaretus) during the eu-and oligotrophication of Lake Constance

About 20 ovaries of the whitefish 'Blaufelchen' ( Coregonus lavaretus L.) of Lake Constance (Bodensee) were collected annually from 1964-1991. Absolute and relative fecundity peaked in the early 1980s, lagging about 2 years behind the maximum of total-P in the water (during the circulation period). Out of the biological time-series of Lake Constance, whitefish fecundity is the only one known to follow the phosphorus curve, and this may be the first time-series documented case of parallel trend reversal in the trophic state of a lake and fish biology. It is concluded that fish fecundity serves, at best, as an unspecific monitor of the overall well-being of the fish.     

Different growth trends of whitefish (Coregonus lavaretus) forms in the northern Baltic Sea

The sea growth of two whitefish forms, anadromous (Coregonus lavaretus lavaretus) and sea‐spawning (Coregonus lavaretus widegreni), was analysed using samples collected from the commercial sea catch in the Gulf of Bothnia (GoB) in the northern Baltic Sea during 1998–2014. In the GoB area, these two forms are possible to identify because the gill‐raker number and size at maturity vary between forms. The growth rate of the forms is linked to their feeding area. Sea‐spawning whitefish, which has a feeding migration near its home site, was shorter in the northern GoB (66°N–64°N) at the ages of 3–11 than those in the southern GoB (64°N–60°30′N). In the data, most whitefish were caught with gill nets in the GoB. The mesh sizes of gill nets capturing the anadromous form were mostly 35–45 mm, while those capturing the sea‐spawning form were <35 mm in the northern GoB. It is likely that the different growth trends for small and large whitefish were connected with differences in their recruitment for fishing. The length of anadromous females at the age of four sea years increased significantly, but the length of six‐year‐old anadromous female whitefish decreased over the catch years from 1998–2014. In contrast, the length of slow‐growing sea‐spawning whitefish of six years or older increased significantly in relation to the catch year in the gill‐net catch. The increase in the growth of young age groups in both forms was probably associated with the increasing temperature and the low fishing pressure on small fish. The decreasing age at capture for both forms and the depression of the mean size of old anadromous whitefish are signs of high fishing pressure with a high gill‐net effort that selectively removes the largest and oldest individuals of both forms.     

Effects of freshwater to seawater transfer on osmoregulation,acid-base balance and respiration in river migrating whitefish (Coregonus lavaretus)

Osmoregulation, acid-base balance and respiratory parameters were investigated in whitefish following transfer from freshwater to salt water. Whitefish acclimated successfully to 25 ppt brackish water but died after direct transfer to 32 ppt sea water. Transfer to brackish water induced rapid (<6 h) and permanent increases in plasma [Na⁺], [Cl^–], total [Ca] and [Mg]. The extracellular hyperosmolality effected a transient (<3 days) muscle tissue dehydration and red blood cell shrinkage. Exposure to brackish water decreased both the arterial O₂ tension and whole body O₂ uptake. The extracellular acid-base status changed from an initial respiratory acidosis at 1 h towards a pronounced metabolic acidosis at 48 h of brackish water exposure. Red cell pH_i decreased in parallel with extracellular pH_e, but the in vivo pH_i/pH_e was only 0.26, suggesting some selective protection of red cell pH_i. Plasma cortisol concentration and gill Na⁺, K⁺-ATPase activity increased after exposure to high ambient salinity, reflecting the induction of hypo-osmoregulatory mechanisms. The physiological changes in whitefish are discussed in relation to salinity-induced effects in other salmonid fishes.Abbreviations CO₂ solubility in plasma - water O₂ capacitance coefficient - BW brackish water - C _T total CO₂ content in plasma - FW fresh water - Hb hemoglobin - Hct hematocrit - M _b body mass of fish - MCHC mean cellular hemoglobin concentration - PCO₂ carbon dioxide tension - pH _e extracellular pH - pH _i intracellular pH - PO_{2 in} oxygen tension in water flowing in - PO_{2 out} oxygen tension in water flowing out - ppt parts per thousand - RBC red blood cell(s) - SW sea water - V _m water flows through chamber - OV ₂ ml O₂ consumed per kg per hour     

Human‐induced changes in the reproductive traits of Lake Constance common whitefish (Coregonus lavaretus)

Size‐selective fishery harvest leads to phenotypic changes in fish reproductive traits. When these changes represent an evolutionary response of a stock, they may have severe consequences for future stock dynamics and yields. In freshwater ecosystems, reproductive traits may also be affected by other human impacts such as changes in system productivity. The present study uses regression analysis to evaluate the impacts of changes in lake trophy and of an intensive size‐selective harvest over several decades on the reproductive traits of common whitefish in Lake Constance between 1963 and 1999. Fecundity was strongly linked to lake trophy but was also related to the calendar year, suggesting an evolutionary response to size‐selective harvest and to massive stocking of the lake with hatchery‐reared larvae. The present study is an example of how fish reproductive traits are influenced by the combined action of various human impacts: changes in system productivity, size‐selective harvest and massive stocking.     

Limits for the detection of daily otolith increments in whitefish (Coregonus lavaretus L.) larvae

The formation of daily otolith increments in European lake white-fish was evaluated in the laboratory during 40 days at 4, 6, and 8 °C under limited and ad libitum food supply. Daily increments were easily identified in larvae reared at 8 °C. At 6 °C and, more, at 4 °C, an unstructured perinuclear zone was formed, and daily increments could only be recognized in the light microscope starting from 10 to 35 days after hatching. In some larvae from the 4 °C groups, no increments at all were found. Scanning electron microscopy (SEM) could not improve increment resolution. Only those increments could be viewed by SEM which were visible in the light microscope as well. We conclude that whitefish larvae which experience low (4–6 °C) temperatures during their first weeks of life, hence those which hatch in the lake, cannot be aged by the currently employed preparation techniques.     

Parallel and non‐parallel morphological divergence among foraging specialists in European whitefish (Coregonus lavaretus)

Parallel phenotypic evolution occurs when independent populations evolve similar traits in response to similar selective regimes. However, populations inhabiting similar environments also frequently show some phenotypic differences that result from non‐parallel evolution. In this study, we quantified the relative importance of parallel evolution to similar foraging regimes and non‐parallel lake‐specific effects on morphological variation in European whitefish (Coregonus lavaretus). We found evidence for both lake‐specific morphological characteristics and parallel morphological divergence between whitefish specializing in feeding on profundal and littoral resources in three separate lakes. Foraging specialists expressed similar phenotypes in different lakes in both overall body shape and selected measured morphological traits. The morphology of the two whitefish specialists resembled that predicted from other fish species, supporting the conclusion of an adaptive significance of the observed morphological characteristics. Our results indicate that divergent natural selection resulting from foraging specialization is driving and/or maintaining the observed parallel morphological divergence. Whitefish in this study may represent an early stage of divergence towards the evolution of specialized morphs.     

Discrete foraging niches promote ecological, phenotypic, and genetic divergence in sympatric whitefish (Coregonus lavaretus)

Natural populations often vary in their degree of ecological, morphological and genetic divergence. This variation can be arranged along an ecological speciation continuum of increasingly discrete variation, with high inter-individual variation at one end and well defined species in the other. In postglacial fishes, evolutionary divergence has commonly resulted in the co-occurrence of a pelagic and a benthic specialist. We studied three replicate lakes supporting sympatric pelagic and benthic European whitefish (Coregonus lavaretus (L.)) morphs in search for early signs of possible further divergence into more specialized niches. Using stomach content data (recent diet) and stable isotope analyses (time-integrated measure of trophic niche use), we observed a split in the trophic niche within the benthic whitefish morph, with individuals specializing on either littoral or profundal resources. This divergence in resource use was accompanied by small but significant differences in an adaptive morphological trait (gill raker number) and significant genetic differences between fish exploiting littoral and profundal habitats and foraging resources. The same pattern of parallel divergence was found in all three lakes, suggesting similar natural selection pressures driving and/or maintaining the divergence. The two levels of divergence (a clear and robust benthic – pelagic and a more subtle littoral – profundal divergence) observed in this study apparently represent different stages in the process of ecological speciation.     

A proteomics approach reveals divergent molecular responses to salinity in populations of European whitefish (Coregonus lavaretus)

Osmoregulation is a vital physiological function for fish, as it helps maintain a stable intracellular concentration of ions in environments of variable salinities. We focused on a primarily freshwater species, the European whitefish (Coregonus lavaretus), to investigate the molecular mechanisms underlying salinity tolerance and examine whether these mechanisms differ between genetically similar populations that spawn in freshwater vs. brackishwater environments. A common garden experiment involving 27 families in two populations and five salinity treatments together with a large-scale, high-resolution mass spectrometry experiment that quantified 1500 proteins was conducted to assess phenotypic and proteomic responses during early development, from fertilization until hatching, in the studied populations. The populations displayed drastically different phenotypic and proteomic responses to salinity. Freshwater-spawning whitefish showed a significantly higher mortality rate in higher salinity treatments. Calcium, an ion involved in osmotic stress sensing, had a central role in the observed proteomic responses. Brackishwater-spawning fish were capable of viable osmoregulation, which was modulated by cortisol, an important seawater-adaptation hormone in teleost fish. Several proteins were identified to play key roles in osmoregulation, most importantly a highly conserved cytokine, tumour necrosis factor, whereas calcium receptor activities were associated with salinity adaptation. These results imply that individuals from these populations are most likely adapted to their local environments, even though the baseline level of genetic divergence between them is low (F(ST)=0.049). They also provide clues for choosing candidate loci for studying the molecular basis of salinity adaptation in other species. Further, our approach provides an example of how proteomic methods can be successfully used to obtain novel insights into the molecular mechanisms behind adaptation in non-model organism.     

Helminth assemblages of whitefish (Coregonus lavaretus) in interconnected lakes: similarity as a function of species specific parasites and geographical separation

This article examined the composition of parasite assemblages of whitefish (Coregonus lavaretus) in 8 interconnected lakes in northeastern Finland and evaluated the role of coregonid specific parasites and the geographical distance between populations in determining the similarity of the assemblages. Parasite assemblages were compared using the Jaccard qualitative similarity index and a quantitative similarity index and by incorporating the allogenic-autogenic species concept and the effects of 2 corresponding measures of geographical distance between the lakes. The majority of the parasite species found (10 of 14) were specific to salmonids. Similarity of assemblages of autogenic parasites between the lakes was negatively correlated with geographical distance. The dominance of 2 parasite species, the whitefish specialist Ichthyocotylurus erraticus and the generalist Ergasilus sieboldi, was also demonstrated. We concluded that the high proportion of widespread parasite species specific to coregonids is an important determinant of similarity in these assemblages. However, ecological factors were likely to contribute to qualitative (presence of species) and quantitative (difference in abundances) differences between lakes, in the case of autogenic parasites, their importance increasing with geographical separation.     

Seasonal changes in the habitat use and movements of adult European grayling in a large subarctic river

In late summer (13 August–13 September 1998), at water temperatures of 12·0–15·7° C, grayling ( n =14) stayed mainly in the riffle-section where they were captured in a large regulated river in northern Finland, moving little between consecutive days. In autumn (2–30 October 1998), at 1·7–6·7° C, the fish ( n =16) migrated to potential overwintering sites 0–14 km up- or downstream by mid October, moving mainly short distances thereafter. The daily movement rates, and the total ranges covered by the fish in late summer and autumn were 54±32 m (mean± s.d ) and 1053±1636 m, and 190±168 m and 3135±1850 m, respectively. In autumn the fish used deeper habitats (most suitable range 150–400 cm) with lower current velocities (20–80 cm s⁻¹) and finer bottom substrata (mainly sand) than in late summer (depth 100–325 cm, velocity 30–110 cm s⁻¹, and cobble-boulder substrata).     

Evolutionary history of the European whitefish Coregonus lavaretus (L.) species complex as inferred from mtDNA phylogeography and gill-raker numbers

We compared mitochondrial DNA and gill-raker number variation in populations of the European whitefish Coregonus lavaretus (L.) species complex to illuminate their evolutionary history, and discuss mechanisms behind diversification. Using single-strand conformation polymorphism (SSCP) and sequencing 528 bp of combined parts of the cytochrome oxidase b (cyt b) and NADH dehydrogenase subunit 3 (ND3) mithochondrial DNA (mtDNA) regions, we documented phylogeographic relationships among populations and phylogeny of mtDNA haplotypes. Demographic events behind geographical distribution of haplotypes were inferred using nested clade analysis (NCA) and mismatch distribution. Concordance between operational taxonomical groups, based on gill-raker numbers, and mtDNA patterns was tested. Three major mtDNA clades were resolved in Europe: a North European clade from northwest Russia to Denmark, a Siberian clade from the Arctic Sea to southwest Norway, and a South European clade from Denmark to the European Alps, reflecting occupation in different glacial refugia. Demographic events inferred from NCA were isolation by distance, range expansion, and fragmentation. Mismatch analysis suggested that clades which colonized Fennoscandia and the Alps expanded in population size 24 500-5800 years before present, with minute female effective population sizes, implying small founder populations during colonization. Gill-raker counts did not commensurate with hierarchical mtDNA clades, and poorly with haplotypes, suggesting recent origin of gill-raker variation. Whitefish designations based on gill-raker numbers were not associated with ancient clades. Lack of congruence in morphology and evolutionary lineages implies that the taxonomy of this species complex should be reconsidered.     

Reactions in individual fish to strobe light. Field and aquarium experiments performed on whitefish (Coregonus lavaretus)

This study describes how individual whitefish Coregonus lavaretusreact to strobe light. Field experiments were performed in a net enclosure on fish tagged with ultrasonic transmitters. A strobe light array was switched on near the tagged fish. The fish moved away from the light and increased their swimming speed. Aquarium experiments under controlled conditions were carried out in rearing tanks at Saimaa Fisheries and Aquaculture Station in Finland. A strobe light was directed from the side of the basin just ahead of, directly at, and behind the fish at a close range. In the first two cases fish responded by a distinct turn and a change in swimming direction away from the light. The fish did not change its swimming direction when light was aimed from behind. It is concluded that strobe light may be used to prevent fish from swimming into a specific area. Implications for development of new fishing equipment and research concerning fishes in areas with water power stations is briefly discussed.     

Relationship between annual variation in reservoir conditions and year-class strength of peled (Coregonus peled) and whitefish (C. lavaretus)

We estimated year-class strength (YCS) of whitefish and peled by VPA in two boreal reservoirs Lokka and Porttipahta with mean regulation of water level amplitudes of 2.34 and 2.49 m, respectively. YCS of peled Coregonus peled(Gmelin) was related to winter drawdown most probably via egg desiccation. In the relatively shallow Lokka dissolved oxygen deficiency in late winter also had a negative impact on survival of eggs and year-class strength. In contrast to peled, whitefish Coregonus lavaretus(L.) indicated no significant relationship with winter drawdown or dissolved oxygen level. The obvious reason for the pattern is that peled spawn on the reservoir bottom, but whitefish spawn in the ascending rivers, which are mainly unaffected by reservoir regulation.     

Site fidelity and activity patterns of a humphead wrasse, Cheilinus undulatus (Labridae), as determined by acoustic telemetry

This study examines the short-term site fidelity, the home range size and the activity patterns of one specimen of the emblematic and threatened humphead wrasse, Chelinus undulatus. Four omnidirectional hydrophones were deployed for 7 months in the mooring area of Larégnère marine reserve (New Caledonia). The hydrophone network covered 50 ha. The monitored fish was caught in the study site and surgically implanted with an ultrasonic transmitter. The fish displayed a regular diurnal rhythm during 25 days before leaving the area. Different patterns of spatial use were identified. The temporal succession of these patterns indicates that the minimum distance covered by the fish every day ranged between 20 and 200 m. Its home range was at least 5 ha. A decrease of the detection was observed before the departure of the fish. This decrease coincided with the beginning of the reproduction period in New Caledonia and supports the hypothesis of a fish departure for the reproductive event. Despite the survey of only one specimen, the home range and movement calculated using acoustic telemetry are different in scale than the reported scuba-based visual estimations. Such information is essential for the development of management policies and conservation measures of large reef fish species highly vulnerable to disturbance and fishing pressure.     

Radioactive strontium (85Sr) in marking whitefish [Coregonus lavaretus (L.)] larvae and the dispersal of larvae from river to sea

Anadromous whitefish [ Coregonus lavaretus (L.)] yolk-sac larvae were marked with a radioactive strontium isotope, ⁸⁵Sr. Before marking the method was tested in accumulation-elimination tests using various strontium concentrations. The first-phase effective half-life of ⁸⁵Sr in yolk-sac larvae was 4 days. The mean ⁸⁵Sr radioactivity was 81 Bq per larva at the day of release and the half-life of the slow component was 51 days, so the strontium label was easily detected up to 60 days after marking. The larvae were released into the River Simojoki about 12 km upstream from the river mouth and recaptured from early May to the end of July. Marked larvae were found at the river mouth 12–13 days after release, while the last marked fish in the river were found at 16 days. It is likely that all leave the river within 2 to 3 weeks post-hatching. Two months after marking some larvae had attained a distance of 10 km from the river mouth.     

Seasonal movement of the American horseshoe crab Limulus polyphemus in a semi-enclosed bay on Cape Cod,Massachusetts (USA) as determined by acoustic telemetry

<正> American horseshoe crabs Limulus polyphemus were tracked using acoustic telemetry and traditional tagging in asemi-enclosed bay on Cape Cod (Pleasant Bay),Massachusetts,USA,to determine seasonal movement patterns.Fifty-five activelyspawning females were fitted with transmitters in 2008 and 2009 and were tracked using acoustic telemetry from May 2008through July 2010.Fifteen crabs with transmitters also had archive depth-temperature tags attached.In addition,over 2000spawning crabs (males and females) were tagged with US Fish and Wildlife (USFWS) button tags over the same period.Ninety-one percent of the crabs with transmitters were detected during this study.In the spring,crabs were primarily located inthe northern section of the bay near spawning beaches,whereas in the fall crabs moved towards the deeper portions of the bay,and some may have overwintered in the bay.There was evidence that a majority (58%-71%) of the females with transmittersspawned in two sequential seasons.One archive tag was recovered resulting in a year-long continuous record of depth and temperaturedata that,when integrated with telemetry data,indicated that the crab overwintered in the bay.The live recapture rate ofcrabs with USFWS button tags was 11%,with all re-sighted crabs except one observed inside Pleasant Bay.Eighty-three percentof recaptures were found within 2.5km of the tagging location,and 51% were observed at the same beach where they were tagged.This study provides further evidence that horseshoe crabs in Pleasant Bay may be philopatric to this embayment     

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

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