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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Predation of perch on vendace larvae: diet composition in an oligotrophic lake and digestion time of the larvae

The diet of perch Perca fluviatilis was studied to reveal possible predation on vendace Coregonus albula larvae in an oligotrophic lake. Perch diet changed with the size of the fish: small perch ate mainly zooplankton and the diet shifted more to benthic invertebrates and fishes in larger perch. There were also annual and spatial differences in the diet, probably reflecting differences in the availability of prey animals. Perch predation on vendace larvae was only observed in the area with high availability of the larvae. The result suggested strengthened predation when the density of the larvae increases. According to bioenergetics modelling, the perch population increased natural mortality of vendace larvae only marginally. Food intake of spawning female perch was slightly reduced, whereas spawning males fed similarly to non-spawning males. Hence, the spawning period of perch was only a minor refuge for vendace larvae. Laboratory experiments of perch digestion rate demonstrated that, due to rapid digestion of the small fish larvae, diet sampling interval should not be >2 h in the field.     

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.     

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.     

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.     

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 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.     

Egg quality and female condition in vendace (Coregonus albula L.) before and during spawning

The fat content of muscle, liver and eggs of vendace (Coregonus albula L.) were measured during the maturation of ovaries in summer. The fat content of eggs was at its highest in July and that of muscle in August during the optimal feeding season. Significant differences were not found in fat content between lakes Suvasvesi and Southern-Kallavesi in central Finland. There were no essential differences in fatty acid composition between muscle, liver and eggs of Lake Suvasvesi vendace. During the last month before spawning in October the gross energy and fat contents of eggs decreased in Lake Suvasvesi vendace. The fat and vitamin C contents of muscle as well as the liver index of females decreased during this time also. In males the corresponding decrease did not occur, indicating that spawning is a greater stress for females than for males. The egg quality and condition of Lake Inarijärvi vendace were about the same as those of Lake Suvasvesi vendace. The unfavourable conditions of Lake Inarijärvi, located in northern Finland, are reflected in the size of ovaries, which is known to be smaller (gonadosomatic index < 20%) than in Lake Suvasvesi (GSI 26–27%). Vendace, then, maintain the quality of eggs at the expense of their number.     

Species introduction promotes hybridization and introgression in Coregonus: is there sign of selection against hybrids?

Species introductions are considered one of the major drivers of biodiversity loss via ecological interactions and genetic admixture with local fauna. We examined two well-recognized fish species, native whitefish (Coregonus lavaretus) and introduced vendace (Coregonus albula), as well as their morphological hybrids in a single lake to test for selection against hybrids and backcrosses in the wild. A representative random subsample of 693 individuals (27.8%) was taken from the total catch of coregonids. This subsample was examined with the aim to select c. 50 individuals of pure whitefish (n = 52), pure vendace (n = 55) and putative hybrid (n = 19) for genetic analyses. The subsequent microsatellites and mitochondrial (mt) DNA analyses provided compelling evidence of hybridization and introgression. Of the 126 fish examined, four were found to be F(1) , 14 backcrosses to whitefish and seven backcrosses to vendace. The estimates of historical gene flow suggested higher rates from introduced vendace into native whitefish than vice versa, whereas estimates of contemporary gene flow were equal. Mitochondrial introgression was skewed, with 18 backcrosses having vendace mtDNA and only three with whitefish mtDNA. Hybrids and backcrosses had intermediate morphology and niche utilization compared with parental species. No evidence of selection against hybrids or backcrosses was apparent, as both hybrid and backcross growth rates and fecundities were high. Hybrids (F(1) ) were only detected in 2 year-classes, suggesting temporal variability in mating between vendace and whitefish. However, our data show that hybrids reached sexual maturity and reproduced actively, with backcrosses recorded from six consecutive year-classes, whereas no F(2) individuals were found. The results indicate widespread introgression, as 10.8% of coregonids were estimated to be backcrosses.     

Dynamics of population of vendace <Emphasis Type="Italic">Coregonus albula</Emphasis> in lake Syamozero (Republic of Karelia)

This work describes the dynamics of the vendace population in Syamozero and provides an evaluation of its current state based on data collected for more than 75 years. The main biological parameters of the fish (body length and mass, gender, maturity stage, and fecundity) are analyzed. The causes of variations in the vendace number from year to year are considered.     

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.     

Successional features of the dynamics of abundance and population structure of the cestode Proteocephalus longicollis (Zeder, 1800) (Cestoda: Proteocephalidae)

Dynamics of abundance and changes of populations structure were investigated in a host-specific parasite of salmonids, cestode Proteocephalus longicollis from the vendace Coregonus albula L. of the Syamozero Lake during the period of more than forty years. A relative stability of the values of infestation indices independently of significant changes in the abundance and population structure of the vendace is established. Changes in the distribution of abundance and population structure of P. longicollis are shown to be connected with the change of the water body's status resulting from eutrophication. These changes can be considered as reliable indices of the succession in freshwater bodies.