Long-term changes in the littoral benthos of a Norwegian subalpine lake following the introduction of the European minnow (<Emphasis Type="Italic">Phoxinus phoxinus</Emphasis>)

The littoral benthos of the subalpine lake, Øvre Heimdalsvatn, has been documented in a series of investigations carried out in 1972, 1976, 1985 and 2000. During this 28-year period there have been major changes in the benthos of the lake following the introduction of European minnow (Phoxinus phoxinus) into the lake where brown trout (Salmo trutta) was formerly the sole species. In 1972 Ephemeroptera, Trichoptera, Plecoptera and Gammarus lacustris dominated the macrobenthos, constituting 85% of faunal numbers, while Chironomidae and Oligochaeta made up only c. 6%. However, by 1976, chironomids and oligochaetes had increased in relative abundance, while G. lacustris declined. This trend towards a dominance of chironomids and oligochaetes was confirmed in 1985 and 2000, although absolute numbers of Ephemeroptera, Plecoptera and Trichoptera increased in 2000 relative to 1972 values. Gammarus lacustris had a 2-year life cycle in Øvre Heimdalsvatn. In 1972 there were significantly more females than males, but by 1976 and through to 2000 there were greater numbers of males. Despite this reduction in females, numbers of juveniles increased, although mortality, probably due to increased predation from minnows, was higher than earlier. The introduction of the alien species, the European minnow, into Øvre Heimdalsvatn has clearly changed the composition and structure of the littoral macroinvertebrate benthos.     

The brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>) in the lake, Øvre Heimdalsvatn: long-term changes in population dynamics due to exploitation and the invasive species,European minnow (<Emphasis Type="Italic">Phoxinus phoxinus</Emphasis>)

Studies of the brown trout (Salmo trutta) population in the Norwegian subalpine lake, Øvre Heimdalsvatn, over a 50-year period have revealed major changes in population dynamics. In 1958, the population density was high, with individuals stagnating in growth at lengths below 30 cm. After heavy exploitation during the years 1958–1969, the number of older fish declined substantially, and growth rates increased significantly. Since 1969, the European minnow (Phoxinus phoxinus) have been observed in the lake, with increasing densities from 1977–1978 to 1999–2000. The age structure of the brown trout population has changed markedly from the 1960s to the period 1993–2006. Annual recruitment significantly declined, from an average number of 3746 individuals in age-class 4 during the period 1958–1966 to an average of 1668 during the period 1993–2006. However, due to lower exploitation rates, the number of old fish was significantly higher in the latter period. The summer diet of brown trout has changed substantially from a dominance of the large crustaceans Lepidurus arcticus and Gammarus lacustris to a high occurrence of European minnows, while L. arcticus has become practically absent from the diet. There was a negative relationship between brown trout biomass and annual length increment. However, despite a brown trout biomass at the same level during the years 1993–2006 as in the 1960s, annual individual growth rates have significantly declined. The reduced recruitment and reduced annual growth rates of the brown trout, as well as changes in the diet, are most likely related to the introduction and establishment of the invasive species, the European minnow.     

Between-year variations in the development of crustacean zooplankton in the Norwegian subalpine lake, Øvre Heimdalsvatn

Data from the International Biological Programme (IBP) and subsequent studies have been re-analysed to test the two hypotheses which previously have been suggested concerning the zooplankton in the mountain lake, Øvre Heimdalsvatn: (1) the average temperature in June, more than other summer months, is affecting the growth rate and population densities of zooplankton in the lake, (2) the invasion of the European minnow (Phoxinus phoxinus) has caused changes in the zooplankton community. The analyses have demonstrated that the June temperature strongly affects the growth rate of all the zooplankton species, but that there is no relationship with the population maxima. The species composition in the crustacean zooplankton has not changed between 1969 and 1999, and any direct impact of the minnows on the zooplankton community could not be detected. Indirectly, the minnows may have reduced the density of invertebrate predators, and thus caused an increase in juvenile survival and increased summer maximum density of Bosmina longispina. The variation in density of the copepod, Cyclops scutifer, was correlated with the density of Heterocope saliens, most likely the result of predator–prey interactions.     

The Norwegian reference lake ecosystem, Øvre Heimdalsvatn

The subalpine lake, Øvre Heimdalsvatn and its catchment, situated in the mountains of southern Norway, has been the subject of extensive ecosystem studies over the last 50 years covering a wide range of disciplines. The lake is located at 1,088 m a.s.l., has an area of 0.78 km², a maximum depth of 13 m, and a catchment area of 23.6 km². It lies more or less on the limit of the birch forest, while the catchment extends up over 1,800 m a.s.l. and into the high alpine zone. The valley is little influenced by human activity and there is no permanent habitation, although the area is used for grazing of domestic livestock during summer. Otherwise the lake and its catchment have always been important for fish and game. The catchment also received major radionuclide fallout from the Chernobyl accident in 1986. The studies include Quaternary history, the development of algorithms for remote sensing of snow conditions, long-term changes in lake ice cover and temperature, the zooplankton and benthic communities, as well as the uptake of Chernobyl radionuclides and mercury by the biota. The consequences of the introduction of the European minnow (Phoxinus phoxinus) on the benthos, zooplankton and the brown trout (Salmo trutta) have also been investigated. The lake and its catchment are widely used in university teaching and thesis work. On account of the substantial knowledge base, Øvre Heimdalsvatn is well suited to long-term monitoring.     

Predation by brown trout: a major mortality factor for sexually mature European minnows

Brown trout Salmo trutta in the subalpine lake, Øvre Heimdalsvatn, showed large temporal variation in the rate of predation on the introduced minnow Phoxinus phoxinus population. Minnows were found in the stomachs of brown trout between 16 and 38 cm L _T. Significantly greater predation was recorded shortly after ice break at the end of June 1999, with frequencies of 9 and 20% within the L _T classes 16–29·9 cm and  ≥30 cm, respectively. Predation on minnows was only occasionally detected during July, August and September. The high level of predation coincided with minnow spawning, and lengths of consumed minnows were equal to those of sexually mature individuals. Accepting a causal link between minnow spawning, which lasted c . 3 weeks, and the contemporary high rate of predation, the estimated annual consumption of minnows by the brown trout population would be 138 kg wet mass. Although most of the annual consumption of minnows by brown trout (90%) occurred within a very short period (3 weeks), it accounted for a significant proportion (60%) of the annual loss in biomass of the sexually mature part of the population.     

Long-term trends and variation in <Superscript>137</Superscript>Cs activity concentrations in brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>) from Øvre Heimdalsvatn,a Norwegian subalpine lake

The Norwegian mountain lake, Øvre Heimdalsvatn, received major fallout from the Chernobyl accident in the spring of 1986. Activity concentrations of ¹³⁷Cs have been monitored annually in the brown trout (Salmo trutta) population of the lake over a 22-year period since 1986. The activity concentrations of ¹³⁷Cs in brown trout have declined over the period, although in recent years there has been little or no reduction in activity concentrations. Throughout the period, there has been considerable individual variation in ¹³⁷Cs activity concentrations. Within a single year, fish weight was the most significant factor affecting activity concentrations of ¹³⁷Cs in individual fish, although sex and age·weight were significant explanatory variables. The “size effect” was not significant during the initial period after fallout and in some recent years. During the first years after fallout, the ecological half-life of ¹³⁷Cs in brown trout was 3–4 years, but during the last decade this has increased significantly and has approached the physical half-life of 30 years for ¹³⁷Cs, suggesting a dynamic equilibrium between catchments inputs, possible remobilisation from lake sediments and lake outputs of ¹³⁷Cs.     

Biology of the minnow Phoxinus phoxinus and its interactions with brown trout Salmo trutta in Øvre Heimdalsvatn, Norway

Minnows have recently been introduced into the mountain lakes of southern Norway. In Øvre Heimdalsvatn, an intensively studied lake, the minnow population has increased rapidly during the last decade. Studies were performed on feeding habits, habitat preferences, age, growth, energy content, densities, parasites, predation and spawning activities. Several aspects were found to be different from other studies, e.g. higher age of sexual maturation, lower growth rate, no feeding on plants, and low infection of parasites. The interaction with the only other fish, the brown trout, is discussed in relation to the minnow population. In 1978 the minnows constituted little more than 10% of production and food consumption of the trout population, and only limited food competition and predation was documented. Increasing competition is likely with a greater minnow population.     

Fishing-induced changes in predation pressure by perch (<Emphasis Type="Italic">Perca fluviatilis</Emphasis>) regulate littoral benthic macroinvertebrate biomass,density, and community structure

We aimed to study whether the varying changes in predation pressure by perch (Perca fluviatilis) reflect the biomass, density, and community structure of the benthic macroinvertebrates. Prey preference is size-dependent, and overall predation pressure is density dependent, and thus the size structure of the P. fluviatilis population should affect the structure of the macroinvertebrate community, and the population density of P. fluviatilis should reflect the overall density of benthic macroinvertebrates. We sampled the littoral benthic community in a boreal lake that had been divided into two parts that were subjected to two different fishing procedures during 2007–2012 period and analyzed the macroinvertebrate diet of fish. The benthic macroinvertebrate community reflected the predation pressure. Total macroinvertebrate biomass increased during the study period in the lake division with a non-size-selective fishing procedure (NSF), i.e., all invertivorous perch size-classes targeted, but decreased in the section with negatively size-selective fishing procedure (SSF), i.e., large invertivorous individuals ≥ 16 cm were not targeted. This difference was a result of the increase in large-sized species, such as Odonata, for the NSF procedure and decrease in the SSF procedure. In contrast to total biomass, total macroinvertebrate density did not show a response to predator size structure but rather total macroinvertebrate density decreased with increasing fish density. The study demonstrates the effect of predation pressure of P. fluviatilis on benthic communities, thus highlighting the keystone predator role of the species in boreal lakes and gives more insight on the multiple effects of fish predation on littoral benthic communities.     

On the diversity of fishes in mountain lakes of the Amur basin

Results of the study of 11 lakes situated in the mountainous part of the Amur River basin are reported. Six fish species, namely, blunt-snouted lenok Brachymystax tumensis, Baikal-Lena grayling Thymallus baicalolenensis, Siberian stone loach Barbatula toni, Lagowski’s minnow Rhynchocypris lagowskii, Czekanowski’s minnow Rh. czekanowskii, and Amur sculpin Cottus szanaga, were found in five lakes. Four species were found in one lake (Lake Pereval’noe, basin of the Amgun’ River), and two species were found in each of the remaining lakes. Lenok and Amur sculpin were the most common species. The fauna of Lake Okonon (basin of the river Zeya) was the most specific and did not have analogs among the lake faunas; Czekanowski’s minnow and Baikal black grayling were found in this lake. The lenok ecotype that formed in the mountain lakes differed from the river lenok in some regards. The potential pathways of formation and protection of mountain lake ichthyocenoses of the Amur basin are discussed.     

Introduction of the European minnow into a subalpine lake: habitat use and long-term changes in population dynamics

Minnows Phoxinus phoxinus , studied 30 years after the first record of the species in the subalpine Lake Øvre Heimdalsvatn, Norway, ≥55 mm L _T, were estimated to have densities of c . 4.7 kg ha⁻¹ (120 000 fish) in June 1999 and 2.1 kg ha⁻¹ (63 000 fish) in June 2000. The population was characterized by low individual growth, delayed age of maturity and lived longer when compared to values reported in a previous study in the early phase of its establishment, and other values reported in the literature. Most minnows reached sexual maturity at 4–5 years and >55 mm L _T. Although the estimated annual survival of minnows >55 mm was low ( S =0.2), ages up to 13 years were recorded. Despite a moderate increase in the population size during the last 20 years, the present reduction in individual growth, followed by delayed age of maturity, suggested the existence of density-dependent effects on the population dynamics of the minnows. The minnows were restricted to the littoral zone and near bottom areas. A vertical or horizontal expansion in habitat use was probably prevented by the presence of piscivorous brown trout Salmo trutta .     

Long-term change of the littoral Cladocera in the Tatra Mountain lakes through a major acidification event

In our study, we focused on littoral Cladocera living and feeding in shallow shore parts of 46 mountain lakes in the Tatra Mountains (Slovakia and Poland). The studied lakes underwent a major acidification event in the 1980s and are now in the process of recovery. Lakes were divided into three categories based on their sensitivity to acidification: 5 extremely sensitive (ES), 11 acid sensitive (AS), and 30 non-sensitive (NS) lakes. In our study, we included historical data from the literature, and data from sediment core and littoral samples, which together represent the evolution of the littoral communities from a pre-industrial period up to the present. In total, 11 littoral species were found belonging to three cladoceran families. Most of the species were members of the family Chydoridae: Alona affinis, A. quadrangularis, A. rectangula, A. guttata, Acroperus harpae, Alonella excisa, A. nana, Chydorus sphaericus, and Eurycercus lamellatus. One species belonged to each family Daphniidae (Ceriodaphnia quadrangula) and Polyphemidae (Polyphemus pediculus). The most numerous littoral taxa were Alona affinis, Acroperus harpae, and Chydorus sphaericus. All species reacted to decreased pH levels during peak acidification in the 1980s by disappearing from most of the lakes of all categories; the only persisting species was Chydorus sphaericus. Most species returned to the lakes when pH started to increase in the 1990s, although their return was noticeably slower in AS lakes. Alona quadrangularis decreased its distribution range over the studied period; Polyphemus pediculus was mostly detected in the 1910s only. The number of species was highest in all lake categories when dwarf pine was present in the lake catchment. On the whole, the littoral community was richest in NS lakes.     

Aquatic oligochaete fauna (Annelida,Clitellata) in Lake Tonle Sap and adjacent waters in Cambodia

In surveys conducted during 2000–2005, 39 taxa of aquatic oligochaetes belonging to the families Enchytraeidae and Naididae were found in Lake Tonle Sap in the Mekong River Basin, Cambodia. Dominated by naidines and pristinines (29 taxa), they mainly comprised widely distributed species as well as South Asian and Southeast Asian species. Among the four areas studied, the littoral regions of the lake—where inundated forests and aquatic vegetation developed during the flooded seasons—presented the highest number of species. Submerged vegetation in the littoral Lake Tonle Sap harbored abundant epiphytic oligochaetes, especially Stylaria fossularis. In contrast to the rich abundance of naidine and pristinine fauna, tubificines and ryhacodrilines were scarce in and around the lake, irrespective of the vegetation in their habitats. Several Aulodrilus species and Branchiura sowerbyi were the main representatives of the benthic oligochaete assemblages throughout the offshore zone of the lake without vegetation. It is noteworthy that the widely distributed tubificines Tubifex tubifex and Limnodrilus hoffmeisteri were not found in any surveyed locality.     

Taste attractiveness of free amino acids and their physicochemical and biological properties (as exemplified by fishes)

Using fishes (32 species, 11 families) as an example, the relationship between the taste attractiveness of free amino acids (L-isomers) and their physicochemical and biological properties was analyzed. It was shown that essential amino acids, most nutritionally required for an organism, have lower taste attractiveness for fishes than nonessential amino acids. Only in 6 of the 32 tested species (sunbleak Leucaspius delineatus, European minnow Phoxinus phoxinus, dace Leuciscus leuciscus, chub Leuciscus cephalus, blue gourami Trichopodus trichopterus, pearl gourami Trichopodus leerii) the relationship between the taste attractiveness and molecular weight of amino acids was supported statistically, being negative in all cases. Only in 2 species, a statistically significant correlation between the taste properties of amino acids and the dissociation constant (K₁) was found, positive in the stone loach Barbatula barbatula and negative in the lake char Salvelinus namaycush. A positive correlation between taste preferences and the magnitude of the isoelectric point (pI) of amino acids was found in one species (roach Rutilus rutilus) and a negative correlation in 2 species (brown trout Salmo trutta and Arctic char Salvelinus alpinus erythrinus). A statistically significant correlation between the taste attractiveness and water solubility of amino acids was revealed in 2 species (chum salmon Oncorhynchus keta and navaga Eleginus nawaga), negative in both cases. The flavor, which stimulates food intake, was found to be more often intrinsic to acidic and polar uncharged than basic and nonpolar amino acids, L- than D-isomers, amino acids with an amino group at the α- than β-position. Amino acids are more attractive than their salts. Aromatic amino acids are much less attractive than S-containing or acyclic amino acids. Thus, in most fish species there is no or weak relationship between the taste attractiveness of free amino acids and many of their physical, chemical and biological properties, suggesting a mediated character of this relationship and/or its poor detectability.     

Polymorphism of smolts of pink salmon <Emphasis Type="Italic">Oncorhynchs gorbuscha</Emphasis> in the Indera River (Kola Peninsula)

Pink salmon introduced into the White Sea started to exploit as spawning grounds middle and upper reaches of the river 20 years after its appearance in the Indera River. As a result of this, the migration pathway of smolts and late smolts appeared in addition to early smolts. The intraspecies polymorphism of smolts is confirmed by differences of early and late smolts by body length and weight, migration dates, food spectrum, and indices of stomach fullness. The food spectra of late juveniles of pink salmon coincide with those of parr of Atlantic salmon Salmo salar and of brown trout S. trutta. Greater abundance of late migrants of pink salmon may cause competition of these species for food.     

A separate lowstand lake at the northern edge of Lake Tanganyika? Evidence from phylogeographic patterns in the cichlid genus <Emphasis Type="Italic">Tropheus</Emphasis>

In Lake Tanganyika, lake level fluctuations were shown to have had a major impact on the evolution of littoral species. Many species are subdivided into arrays of populations, geographical races and sister species, each colonizing a particular section of the shore. Their often limited dispersal abilities promoted geographic isolation and, on the long run, allopatric speciation. With more than 120 distinct populations, the genus Tropheus represents the most spectacular and best-studied example of this phenomenon. The present study aims at the fine-scale reconstruction of the spread of two mitochondrial Tropheus-lineages in the very north of the lake, where two species, T. sp. ‘black’ and T. brichardi, occur. Using mtDNA sequences and AFLP-data, we analyzed samples from 21 localities and found a highly complex conglomerate of introgressed populations formed by the repeated contact of two lineages. Our data suggest repeated cross-lake dispersal of T. sp. ‘black’ haplotypes along the ridge between the West and East Ubwari Fault, supporting an additional persisting lowstand-lake in the Bujumbura subbasin at the very north of the lake and highlighting once more the impact of lake level fluctuations on the genetic structure and evolution of stenotopic rock-dwelling cichlid species.     

Dynamics of species composition of fish and some results of ichthyomonitoring in the middle course of the Oka (Ryazan oblast)

The dynamics of species composition of fish from the middle course of the Oka is considered. A total of 53 species of Cyclostomata and fish was previously recorded; however, only 43 species have been found at the present time. Due to different causes, ten species have disappeared, and ten new species have emerged. The species composition and structure of aboriginal fish population are characterized by relative stability; however, the total abundance of fish during the current decade, as compared to the 1970s, has considerably decreased. The sterlet Acipenser ruthenus, lake minnow Phoxinus precnurus, Volga zander Sander volgensis, and freshwater sculpin Cottus gobio belong to rare fish.     

Evaluating Stocking Efficacy in an Ecosystem Undergoing Oligotrophication

Oligotrophication has negatively affected fisheries production in many freshwater ecosystems and could conceivably reduce the efficacy of stockings used to enhance fisheries. In Lake Michigan, offshore oligotrophication has occurred since the 1970s, owing to reductions in total phosphorus (TP) inputs and nearshore sequestration of TP by nonindigenous dreissenid mussels. We evaluated simultaneous effects of stock enhancement and oligotrophication on salmonine species (Chinook salmon Oncorhynchus tshawytscha, lake trout Salvelinus namaycush, and steelhead O. mykiss) that support valuable recreational fisheries. We employed a novel application of an Ecopath with Ecosim model by conducting a full factorial simulation experiment. Our design included multiple levels of salmonine stocking, consumption by invasive quagga mussels (Dreissena bugensis), and TP that were informed by manager interests. Under all levels of TP and quagga mussel consumption, our results showed that stock enhancement could still increase salmonine biomass, but positive responses were stronger for lake trout and steelhead than Chinook salmon. Simulations showed that quagga mussel consumption has deleterious effects on pelagic-oriented prey fishes and Chinook salmon, which feed almost exclusively on the pelagic-oriented alewife (Alosa pseudoharengus). In summary, results from our simulation experiment suggested that lake trout and steelhead are better suited to the current ecosystem than Chinook salmon, and therefore, stock enhancement provides the highest gains for these two species. Furthermore, simulated biomass of all recreational salmonine species increased with increasing TP, indicating the need for managers to consider how potential future oligotrophication will limit the carrying capacity of salmonine biomass in Lake Michigan.     

Tashkent riffle minnow <Emphasis Type="Italic">Alburnoides oblongus</Emphasis> belongs to the genus <Emphasis Type="Italic">Alburnus</Emphasis> (Osteichthyes: Cyprinidae) as inferred from mitochondrial and nuclear DNA markers

Sequences of mitochondrial (cytochrome b) and nuclear (recombination activating gene 1–RAG1) DNA markers were obtained for two species of the genus Alburnoides, the Taskent riffle minnow A. oblongus Bulgakov 1923 and the Terek spirlin A. gmelini Bogutskaya and Coad 2009. Phylogenetic analysis revealed that A. oblongus belongs to the genus Alburnus.     

Peculiarities of the Structure of and Trends in the Macrobenthos Community of the Ermolinskaya Bay Lagoon Ecosystem,Kandalaksha Bay,White Sea

The species composition and spatial and trophic structures of the macrobenthos communities of the Ermolinskaya Bay were studied, and the main changes that had occurred since previous explorations (the 1960s) were analyzed. The most significant changes in the spatial structure of the macrobenthos community (species diversity, population density and biomass, and similarities of the species’ distribution) were observed with regard to the sea level zone: the most pronounced at the upper littoral, followed by the middle littoral, lower littoral, and sublittoral. The longitudinal gradient of the community structure (from the innermost area seaward) was pronounced less strongly. Altogether, four macrobenthos communities can be identified in Ermolinskaya Bay; they differ in species composition, diversity, population density and biomass, and trophic structure: (1) community of the upper littoral with the dominance of collecting deposit feeders, Hydrobia ulvae, and scrapers, Littorina saxatilis; (2) community of the middle littoral with the prevalence of deposit feeders, Hydrobia ulvae and Macoma balthica, seston feeders, Mya arenaria, and, to a lesser extent, Mytilus edulis; (3) community of the lower littoral with the domination of the filtering organisms, Mytilus edulis, to a lesser extent collecting deposit feeders, Hydrobia ulvae and Macoma balthica, and scrapers, Littorina littorea; and (4) community of the sublittoral, where the leading positions belong to the group of collecting deposit feeders, Macoma balthica, to a lesser extent also Hydrobia ulvae (in the innermost area) and Capitella capitata, Arenicola marina (in the marine part of the bay). The gradual siltation of the bay and the reduction of its connection to the sea have led to the development of a littoral complex of species in the sublittoral, whereas the species typical at the sublittoral in the 1960s are now mainly found at the outlet of the bay.     

Assessing the efficacy of corn-based bait containing antimycin-a to control common carp populations using laboratory and pond experiments

Strategic use of oral toxicants could allow for practical and sustainable control schemes for the invasive common carp (Cyprinus carpio, or ‘carp’) if a toxicant selectively targeted carp and not native species. In this study, we incorporated antimycin-a (ANT-A), a known fish toxicant, into a corn-based bait and conducted a series of experiments to determine its toxicity, leaching rate, and species-specificity. Our results showed that ANT-A was lethal to carp at doses ≥ 4 mg/kg and that the amount of ANT-A that leached out of the bait in 72 h was not lethal to carp or bluegill (Lepomis macrochirus). Species-specificity trials were conducted in 227 L tanks, in which carp were stocked with three native species representing families that occur sympatrically with carp in our study region: the fathead minnow (Pimephales promelas), yellow perch (Perca flavescens) and bluegill. These trials showed high mortality of carp (46%) and fathead minnows (76%) but no significant mortality of perch or bluegill. Finally, a pond study, which used the same species composition except for fathead minnows, resulted in 37% morality among adult carp and no mortality among perch or bluegill. Our results suggest that corn-based bait that contains ANT-A could be used to selectively control carp in ecosystems dominated by percids or centrarchids, such as lakes across the Great Plains ecoregion of North America, where carp are especially problematic.