Unpredictability of fish recruitment: interannual variation in young–of–the–year abundance

In 1989–1998, vendace larvae Coregonus albula were sampled in Finnish lakes following a stratified random sampling design. The abundance of young–of–the–year vendace after the first growing season was estimated using catch–per–unit–effort statistics. The number or total area of nursery places hardly limited the recruitment of vendace in the study lakes. The major proportion of prerecruit mortality of vendace occurred in the larval phase and larval sampling produced significant information on young–of–the–year survival. Although larval abundances and recruitment were clearly associated only in Lake Onkamo, generally high larval abundance was needed to produce high number of recruits.     

Population regulation of vendace (Coregonus albula) in Lake Pyhäjärvi, southwest Finland

Previous time series analysis on vendace population dynamics in Lake Pyhäjärvi, 1971–1990, revealed a 2-year cycle in year-class strength, implying powerful density-dependent regulation. Here we have extended this analysis by using multiple regression models to test whether the recruitment series is influenced by density-independent factors. We chose population size with a lag of 1 year as the density-dependent factor; the density-independent factors were the summer water temperature with a lag of 2 years (temperature sum for June, July and August, indicating the year-class strength of predators) and the temperature-derived length of the larval period of vendace. For the years 1972–1990 the coefficient of determination ( r ²) of this regression model was 0·77. We suggest that the basic mechanism producing a persistent 2-year cycle of vendace in Lake Pyhäjärvi is the asymmetrical food competition between age groups. The abundance of predators in the lake and the warming of the water after the hatching of larvae in spring together determine the final year-class strength of vendace.     

The influence of food competition and host specificity on the transmission of Triaenophorus crassus (Cestoda) and Cystidicola farionis (Nematoda) to Coregonus lavaretus and Coregonus albula (Pisces:Coregonidae) in Finland

As a superior competitor for planktonic food, vendace (Coregonus albula), when abundant, is expected to displace whitefish (Coregonus lavaretus) from feeding on plankton and to force it to rely more on benthic food. The predicted result would be a reduced abundance of the copepod-transmitted cestode Triaenophorus crassus in whitefish, but an increase in the abundance of the nematode Cystidicola farionis transmitted via benthic amphipods. We studied the occurrence of both parasites in whitefish during 1991-1996 in three interconnected areas at Lake Saimaa, Finland, where the densities of the vendace stocks varied due to natural fluctuation in year-class strengths. In accordance with our hypothesis, some indication of the effect of the density of the vendace population on abundance of C. farionis infection in whitefish was found, but not in the case of T. crassus. Only 0.2% of vendace were infected with T. crassus, while up to 100% of the whitefish in the yearly samples harboured the parasite. In further experiments we clarified which copepod species in Lake Saimaa act as first intermediate hosts of T. crassus, and verified from stomach samples the exposure of both whitefish and vendace to those species. Experimental infections indicated that the infectivity of T. crassus is lower for vendace than for whitefish. We suggest that the reason why vendace stock density does not affect T. crassus infection in whitefish is that T. crassus is transmitted in littoral areas during a short period in spring. At that time of the year copepods are abundant and available to both whitefish and vendace, but since the parasite is less infective to vendace. they do not become infected. At other times of the year, dense vendace stocks may force whitefish to shift to benthic food, which includes amphipods transmitting C. farionis. During vendace stock decline, whitefish may, however, continue to feed on plankton and avoid exposure to C. farionis.     

Monthly variability of hydroacoustic fish stock estimates in a deep lake and its correlation to gillnet catches

Monthly (April to November) hydroacoustic surveys and parallel gillnet catches were used to determine vendace Coregonus albula abundance, biomass and population dynamics in a deep oligotrophic lake. By hydroacoustic surveys, recruitment of 0+ year vendace could clearly be detected. In contrast, gillnet catches resulted in low numbers of 0+ year fish, and similarly under-represented proportions of the oldest vendace. Consequently, the correlation between hydroacoustics and gillnets with respect to fish numbers and geometric mean fish total length was high only for the age groups 1 + to 4+ years. Annual variability in hydroacoustic estimates offish abundance and biomass was high (CV=26–29%) which reflects the seasonal population dynamics of vendace.     

On the origin of Solovetskaya vendace <Emphasis Type="Italic">Coregonus albula</Emphasis> and of the syamozero smelt <Emphasis Type="Italic">Osmerus eperlanus</Emphasis>

The genetic analysis is made on the population of the European smelt Osmerus eperlanus occasionally introduced to Lake Syamozero (Karelia) and of the vendace Coregous albula supposedly acclimatized in Solovetskie Islands as a result of fish cultural activities of the Solovetskii Monastery. They are compared with several natural populations—possible donors for both introductions. Genetic variation in a sample of smelt was estimated by means of restrictase analysis of mtDNA (fragment ND1/ND2) in samples of vendace—by means of allozyme analysis of six isoenzyme systems. The probable population for the Syamozero smelt is the population of Onega Lake in spite of the previously noted greater morphological similarity of colonizers with the smelt of Ladoga Lake. A high level of genetic variation in the Syamozero smelt in comparison with native populations indicates that, beside the introduction from Onega Lake, there were repeated introductions from neighboring water bodies. The genetic analysis of the Solovetskaya vendace does not prove that the vendace appeared on islands due to acclimatization. Frequencies of alleles of allozyme loci in the Solovetskaya population significantly differ from frequencies in continental populations. Still, it is compared with some populations of the Arkhangelsk oblast. Estimations of genetic diversity of the Solovetskaya vendace turned out to be comparable with those in native populations.     

Year-class fluctuations in vendace,Coregonus albula (Linnaeus): Who's got the upper hand in intraspecific competition?

The vendace Coregonus albula (L.) populations in the lakes Mjøsa and Osensjøen exhibited fluctuating year-class strength. In Mjøsa, a strong year-class emerged every third year, except for the four year period between the strong year-classes 1969 and 1973. The difference between the strong and weak year-classes decreased from the 1960s, through the 1970s to the 1980s. The Mjøsa vendace matured sexually at age 2 +, and more than ten sexually mature age-groups were present in the population. Growth ceased at maturation, and asymptotic length was 23.6 cm. In Osensjøen, one strong year-class (1969) dominated the population during the period 1976–1987. The Osensjøen vendace matured sexually at age 3, and more than 15 sexually mature age-groups were present in the population. Growth ceased at maturation, and asymptotic length was 28.4 cm. In both lakes, vendace fed on crustacean zooplankton in the epilimnion throughout summer and autumn. Our data indicate that regular year-class oscillations occur as a result of the juvenile survival being negatively correlated to the number of adults.     

Patterns in vendace recruitment in Lake Pyhäjärvi, south-west Finland

A vendace Coregonus albula population has long formed the basis of a flourishing fishery in Lake Pyhäjärvi, south-west Finland. Between 1971 and 1990, it exhibited variations in year-class strength generally characterized by a 2-year cycle of alternating strong and weak year classes, which implied density-dependent regulation probably due to asymmetrical food competition between age groups. The abundance of piscivorous predators in the lake and the wanning of water after the hatching of larvae in spring together determined the final year-class strength of vendace. Since 1990, year-class sizes have remained very low and the 2-year cycle has been disrupted. The continuously low levels of recruitment could not be explained by an increased larval mortality, although the decline started with 2 successive years with very high mortality of this life stage which reduced the whole spawning stock to a fraction of earlier levels, and consequently led to low larval numbers such that between 1988 and 1996 larval abundance was positively correlated with the spawning stock size. The importance of the spawning stock size to recruitment could also be seen in the whole 26-year time series as a significant correlation between year-class sizes with a 2-year lag. The observed recruitment patterns of the vendace stock in Lake Pyhäjärvi thus show the importance of spawning stock size, intraspecific competition, predator effects and abiotic environmental variation.     

Using a 1-D mixing model to assess the potential impact of year-to-year changes in weather on the habitat of vendace (Coregonus albula) in Bassenthwaite Lake, Cumbria

1. Bassenthwaite Lake in Cumbria is one of only two English lakes containing a population of vendace (Coregonus albula). The spatial distribution and survival of this fish is strongly influenced by the temperature and oxygen content of the water. In summer, this fish moves into deeper, colder water but avoids areas where the oxygen content is low. 2. In recent years, there has been a dramatic decline in the number of vendace found in the lake, a trend that may have been exacerbated by a succession of warm summers. Bassenthwaite only becomes stably stratified during calm, warm periods when a significant proportion of the deep water becomes anoxic. 3. Here, a one dimensional (1‐D) process‐based temperature‐oxygen model is used to simulate the year‐to‐year variations in the severity of these ‘extreme events’. The model is validated using field measurements acquired in the 1990s and used to predict the range of depths accessible to the vendace. 4. An empirical, weather‐driven model is then used to ‘hindcast’ the mixing characteristics of the lake in the 1980s and estimate the proportion of the habitat lost during warm, calm summers. These simulations show that periods of stable thermal stratification have become increasingly common in recent years. In the 1980s, only one ‘extreme event’ was identified but four such events were recorded in the 1990s. 5. The results are discussed in relation to the conservation status of the species and the potential effect of climate change on its survival in the English Lake District.     

Fish stock assessments in Lake Konnevesi

In 1977, the average fish yield in the oligotrophic and oligohumic lake Konnevesi was 6.4 kg ha^–1 (2.6 kg vendace, 1.3 kg perch, 0.7 kg whitefish, 0.5 kg roach, 0.3 kg pike and 0.3 kg burbot). In that year the stock of vendace was exceptionally poor. A study was made to determine the optimum strategy for exploiting the stocks. The field work included test fishing (1970–78), sampling of catches (1977–80) and collection of catch per unit effort data (1978–80), and catch statistics (1969, 1970, 1977). The stocks were assessed by the Beverton and Holt method. Whitefish is exploited at a proper level. Fishing of vendace and roach can be increased, and the yield of pike would be greater, if the recruiting age were higher than at present.     

Dispersion of vendace eggs and larvae around potential nursery areas reveals their reproductive strategy

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The use of live and frozen Artemia salina nauplii enriched with fluorochromes for mass‐marking vendace Coregonus albula (L.) larvae

The aim of this study was to investigate the use of Artemia salina nauplii enriched with chosen fluorochromes for mass marking vendace Coregonus albula (L.) larvae. In the experiment, vendace larvae (6 DPH) were fed with live or frozen A. salina nauplii immersed in tetracycline hydrochloride (TC), calcein (Cal) or alizarin red S (ARS) for four subsequent days. More successful effects (marking otoliths) were obtained by feeding the fish with live nauplii enriched with fluorochromes, regardless of the dye type. The highest percentage of marked otoliths (100%) was recorded in the group fed with live nauplii immersed in TC. In the groups fed live or frozen nauplii enriched with Cal and ARS, a lower percentage of marked individuals (63.3%–73.7% and 56.7%–63.3%, respectively) were recorded. The survival rate of vendace larvae in particular groups oscillated between 93.7% and 95.7%. There were no significant differences in the total body length and body weight of the reared vendace larvae among different groups. In conclusion, for mass marking of vendace larvae using a feeding method, fish fed A. salinalive nauplii enriched with TC at a dose of 600 ppm is recommended for fishery practice.     

Minnow predation on vendace larvae: intersection of alternative prey phenologies and size-based vulnerability

Time-intensive sampling was used to study minnow Phoxinus phoxinus density and foraging activity in the littoral area of Lake Lentua at a time of high vendace Coregonus albula larval abundance. Minnow activity and foraging during the late spring-early summer period at low temperatures was found to be mainly nocturnal and quite consistent with features reported in the literature, with the exception of feeding on vendace larvae. The absence of the latter finding from previous studies may be due to previous seasonally limited daytime samplings. The temporal or spatial scale of sampling is decisive when studying foraging on food animals with a brief period of vulnerability and seasonal phenology. However, minnows in Lake Lentua seemed to prefer Bosmina longispina whenever available and low B. longispina density during the first weeks after the break-up of ice directed minnow predation towards the vendace larvae. Estimated gross predation values showed that the minnow has the potential to become a remarkable predator on fish larvae. Slight variations in the vernal timing of the vendace hatching and larval development with respect to minnow activity, both of which are obviously temperature related, may be critical to vendace larval survival in the nearshore zone. However, the predation on the larvae is probably a strong factor only for a short period and the survival of the vendace larvae may be more related to the availability of suitably sized food resources than to predation by the minnow.     

Seasonal variations of lipid fatty acids of boreal freshwater fish species

1. The fatty acid levels of muscle and liver lipids of perch, vendace and rainbow trout (Salmo gairdneri) cultivated in the same area (for comparison) were monitored. 2. The total lipid content in the muscle of perch (Perca fluviatilis) and vendace (Coregonus albula) was less than 50% of that in rainbow trout and a seasonal variation was clear only in vendace. 3. The relative amounts of polyunsaturated fatty acids as well as omega-3 acids were higher in vendace and perch than in cultivated rainbow trouts. Arachidonic acid content was much higher in vendace and perch than in rainbow trout. The content of monoenes was considerably higher in rainbow trout than in free freshwater fish. 4. The seasonal variations in the degree of unsaturation were small in fish muscle. 5. In the muscle of rainbow trout the relative amount of polyunsaturated fatty acids diminished with the increase of total lipids.     

Plasticity in habitat use determines metabolic response of fish to global warming in stratified lakes

We used a coupled lake physics and bioenergetics-based foraging model to evaluate how the plasticity in habitat use modifies the seasonal metabolic response of two sympatric cold-water fishes (vendace and Fontane cisco, Coregonus spp.) under a global warming scenario for the year 2100. In different simulations, the vertically migrating species performed either a plastic strategy (behavioral thermoregulation) by shifting their population depth at night to maintain the temperatures occupied at current in-situ observations, or a fixed strategy (no thermoregulation) by keeping their occupied depths at night but facing modified temperatures. The lake physics model predicted higher temperatures above 20 m and lower temperatures below 20 m in response to warming. Using temperature-zooplankton relationships, the density of zooplankton prey was predicted to increase at the surface, but to decrease in hypolimnetic waters. Simulating the fixed strategy, growth was enhanced only for the deeper-living cisco due to the shift in thermal regime at about 20 m. In contrast, simulating the plastic strategy, individual growth of cisco and young vendace was predicted to increase compared to growth currently observed in the lake. Only growth rates of older vendace are reduced under future global warming scenarios irrespective of the behavioral strategy. However, performing behavioral thermoregulation would drive both species into the same depth layers, and hence will erode vertical microhabitat segregation and intensify inter-specific competition between the coexisting coregonids.     

Resource partitioning between pelagic coregonids in a subarctic watercourse following a biological invasion

If a pelagic ecosystem is invaded by an efficient planktivorous fish, competition theory predicts that habitat and/or diet segregation should occur if the invader and native planktivores are to co-exist. We compared the diet and pelagic habitat use between invading vendace (Coregonus albula) and two native morphs of whitefish (Coregonus lavaretus) in three subarctic lakes located within the same watercourse. No clear vertical habitat segregation was found as vendace prevailed throughout the entire depth range. Zooplankton generally dominated the diet of all three coregonids. No obvious resource partitioning was found in two of the lakes, while in the third lake the diets of whitefish and vendace differed significantly from each other. Here, the predominant pelagic whitefish morph fed extensively on water mites whereas vendace mainly consumed surface insects and to some extent small fish, apparently as a result of highly depleted zooplankton resources. The results suggest that various levels of resource depletion following the vendace invasion caused the observed differences between the lakes, indicating that different levels of resource partitioning were associated with dissimilar competition intensities. At present, coexistence of these congeneric species seems to be possible, although the vendace invasion has apparently resulted in a challenging and unstable situation for the planktivorous whitefish.     

REGULAR PAPERS Habitat distribution and size structure in freshwater fish communities: effects of vendace on interactions between perch and roach

In a study of habitat selection and size structure in three fish species in 115 Swedish lakes, roach Rutilus rutilus used the pelagic habitat to a lesser extent and were lower in pelagic biomass in the presence of vendace Coregonus albula. Size distributions of roach were skewed toward larger sizes in lakes with vendace, although a similar pattern of size-dependent habitat use was found regardless of lake group. In disagreement with predictions, vendace did not affect relative perch Perca fluviatilis biomass, size structure, or the proportion of large perch. However, a higher growth for large perch indicated higher per capita energy intake in lakes with vendace.     

Genetic changes in a vendace Covegonus albula (L.) population, 92 years after introduction

The Osensjmn vendace, Coregonus albula (L.), population was founded in 1895 by introduction of about 90 000 fry from Lake Mjøsa. After 92 years, or about 46 generations of isolation, remarkable genetic changes were evident among the transplants compared to their source population. Electrophoresis of 33 enzyme loci revealed that allele frequencies differed significantly in eight of the 10 polymorphic loci. Nei's unbiased genetic distance was D = 0.012, representing the level of population differentiation usually observed among distant watercourses in vendace. Genetic variability was clearly higher among the transplants: average observed heterozygosity was 14.8% compared with 11.2% among Lake Mjøsa vendace. The high genetic variation among Osensjoen vendace shows that the divergence is not due to a small number of founders or bottlenecks in later generations. Selection by the contrasting environmental conditions between the habitats of the donor and transplanted populations is the most plausible hypothesis to explain the genetic changes in the Osensjsen vendace.     

The genetic relationships of the coregonid fishes of Britain and Ireland indicated by electrophoretic analysis of tissue proteins

The electrophoretic patterns of the general proteins, esterases, creatine kinase and phosphoglucomutase of white muscle from Lough Neagh pollan, Lough Erne pollan, Lake Bala gwyniad, Loch Lomond powan, Haweswater schelly and Bassenthwaite vendace are consistently of three types. It is therefore suggested that the coregonids examined are three distinct species: the pollans; gwyniad, powan and schelly; vendace. Vendace is genetically closer to the gwyniad group than to the pollans.     

The size of a vendace, Coregonus albula L., stock in a deep lake basin monitored by hydroacoustic methods

An acoustic estimate of the number and biomass of a vendace stock (age >2 + years) in a deep basin of Lake Karjalan Pyhäjärvi was made in August 1985. The acoustic data were collected at night during the summer stagnation. The vendace were on the lake bottom during the day and rose by midnight to the hypolimnion; they did not rise to the thermocline or water layers above it. In the research area the mean number of vendace was 1900 fish ha ⁻¹ and the mean biomass was 76 kg ha ⁻¹. Total vendace biomass was 151 in the whole 200-ha research area.     

Populational variability of Proteocephalus longicollis (Cestoda: Proteocephalidae) in the vendace Coregonus albula from lakes of Karelia

Populational variability of Proteocephalus longicollis from the vendace inhabiting various types of Karelian water bodies was studied. High phenotypic diversity of the helminth and polymorphism in the cestode's major functional complexes - attachment, trophic and reproduction - are established. High degree of phenotype pool realization was determined for various P. longicollis populations. Traits belonging to different functional complexes were shown to have different variability patterns. The highest stability was shown for the attachment organs of P. longicollis, which had similar values of qualitative indices and nature of variation. The traits of the trophic and reproduction complexes are more variable. The conclusion was made that the high intrapopulation diversity of P. longicollis is a response to high ecological variability of the vendace, and the distinctions in the phenotype variability of the helminth reflect the differences between population development patterns of the parasite (cestode P. longicollis) and its host (vendace Coregonus albula) in lakes of Karelia.