Diet and movements of pikeperch (Sander lucioperca) in a large oligotrophic lake with an exceptionally high pikeperch yield

Through the recent increases in abundance, pikeperch (Sander lucioperca) has not only become a valuable target for recreational and commercial fishing, but also the most important top predator in many lakes. Generally oligotrophic Lake Höytiäinen (283 km²) sustains one of the most economically valuable pikeperch stocks in Finland, and its annual fisheries yield (up to 5.5 kg/ha) is exceptionally high among Finnish lakes. Here, we followed seasonal movements of fish using conventional tagging, and studied short‐term and long‐term diet of pikeperch by stomach content and stable isotope analyses to examine if the movements were motivated by prey availability. A clear seasonal migration pattern was observed: in summer, 80% of the tag recoveries were obtained from the shallow and mesotrophic northern and western areas, whereas in autumn and winter, the deep and oligotrophic southern area dominated in the recaptures. The diet of pikeperch consisted of the most abundant fish species in the lake, perch (Perca fluviatilis) and roach (Rutilus rutilus), as well as insects. Stable isotope analysis indicated a shift from a dominance of pelagic food sources to littoral food sources during the growing season of pikeperch. Our results suggest that the pikeperch migrations facilitate efficient material transfer from the mesotrophic part while the deeper and oligotrophic part offers a favourable overwintering environment.     

Inter- and size-specific patterns of fish seasonal migration between a shallow lake and its streams

Abstract –  This study used passive telemetry (passive integrated transponders) to evaluate winter migration in three species of cyprinids (roach ( Rutilus rutilus (L.)), white bream ( Blicca bjoerkna (L.)) and rudd ( Scardinius erythrophthalmus (L.))) and their potential predators (pike ( Esox lucius (L.)) and perch ( Perca fluviatilis (L.))) between a shallow lake and its streams. Migration patterns were investigated from October to June, and a substantial part of the roach (40%) and white bream (55%) populations tagged in the lake during autumn migrated during winter into the streams, whereas only very few piscivores (<2%) migrated. In contrast to roach and white bream, only few rudd (<6%) migrated, which is likely a consequence of different overwintering strategies, e.g., rudd overwintering in shallow highly structured habitats. Small rudd migrated more than larger rudd, whereas there were no size-differentiated migration patterns for roach or white bream. Migration of the cyprinid fishes was generally initiated in late October and ended in May, and specific synchronised bursts of migration were observed in December, January and April, suggesting that migration is triggered by one or more proximate environmental cues. The cyprinid fishes generally entered the streams in late afternoon or in the morning, depending on season, but overall migration patterns varied between the three streams. We suggest and discuss that our results have great implications for lake management as well as for the interpretation of seasonal trophic dynamics in shallow lakes.     

The influence of diel period on fish assemblage in the unstructured littoral of reservoirs

Abstract Diel changes in littoral fish assemblage were studied in four reservoirs in the Czech Republic (Central Europe). The sampling was performed by beach seining in an unstructured littoral zone. Perch, Perca fluviatilis L., roach, Rutilus rutilus (L.), bream, Abramis brama (L.), carp, Cyprinus carpio L., ruffe, Gymnocephalus cernua (L.), pikeperch, Sander lucioperca (L.), eel, Anguilla anguilla (L.) and A. brama × R. rutilus hybrids exhibited higher densities at night. Only bleak, Alburnus alburnus (L.) exhibited higher densities during the day. The number of species was higher in night hauls, and mean body size of roach and bleak was also higher at night. Diel changes in fish densities resulted in the change in species composition between day and night. The study has implication for the sampling design to assess littoral fish assemblages using beach seine netting and recommends night sampling for a representative assessment or sampling during both diel periods for a robust assessment.     

Associations of fish with various types of littoral habitats in reservoirs

Fish associations with different types of littoral habitats were studied in four canyon‐shaped reservoirs in the Czech Republic in years 2010 and 2011 by gillnets. Two to three habitats per reservoir–beaches (former meadows), stump fields (former forest) and rubble slopes–were defined and sampled along the longitudinal axis of reservoirs. Effects of reservoir, habitat and locality (position along longitudinal axis) on fish biomass, abundance and species structure were tested for juvenile and adult fish separately. Hierarchical analysis of variance revealed that habitats differed significantly in fish biomass and abundance. Redundancy analysis showed that analysed environmental variables had significant influence on fish community structure. Most variability in community structure was explained by reservoir and then by combination of habitat and slope steepness. Locality position had the smallest influence on community structure. For both adult and juvenile fish total abundance and biomass, the most inhabited habitat was beaches; rubble slopes were the least inhabited. Habitat associations differed among species. Among adults, bream Abramis brama, white bream Blicca bjoerkna and roach Rutilus rutilus were associated with beaches and stump fields, whereas perch Perca fluviatilis, ruffe Gymnocephalus cernuus, asp Aspius aspius and pike Esox lucius were associated with rubble slopes. Bream, white bream, bleak Alburnus alburnus, roach, ruffe and pikeperch Sander lucioperca were associated with beaches among juveniles, whereas the only juvenile associated with rubble slopes was perch. We showed that most common species are associated with distinct habitats and also that utilisation of various littoral habitats differs in general.     

Habitat use of lake trout (Salvelinus namaycush) following species introduction

Abstract – Although introductions of prey species have the potential to significantly alter habitat use by top predatory fish, this aspect has rarely been directly quantified. Introduction of yellow perch (Perca flavescens), a littoral–pelagic prey species, to a small boreal lake previously dominated by littoral cyprinids provided a unique opportunity to examine how a change in forage base influenced habitat use by the sole top predator, lake trout (Salvelinus namaycush). We monitored lake trout pelagic and spatial distribution using acoustic telemetry before (2001) and after (2008) the introduction of perch to determine whether habitat use reflected a deeper, offshore prey community. After accounting for differences in water temperature and dissolved oxygen concentrations between years and the inclusion of a control lake, our data suggest that lake trout habitat use changed after the introduction of yellow perch. Lake trout, on average, were 1.4 m deeper (P < 0.01), reduced their use of littoral habitat by 55% (P = 0.03) and experienced a 71% decrease in home range size (P < 0.01), consistent with a greater offshore habitat overlap between predator and prey after the introduction of yellow perch. This study illustrates how introduced prey species may have a significant influence on habitat use by top predatory fish, while also showing the importance of using direct measurements to quantify behavioural changes.     

The effect of depth, distance from dam and habitat on spatial distribution of fish in an artificial reservoir

Abstract –  Spatial distribution of the fish community in the deep canyon-shaped Římov Reservoir, Czech Republic, was studied using overnight gillnetting fishing in 1999–2007. Effects of depth, distance from the dam to the tributary and habitat type (benthic and pelagic) on fish community structure, catch per unit of effort (CPUE), biomass per unit of effort (BPUE) and average weight were tested. Fish were recorded in all sampled depths and parts of the reservoir. Redundancy analysis revealed that effects of three environmental variables were significant and most variability was explained by depth, then by distance from the dam and habitat type. CPUE and BPUE of all species decreased with depth and responses of individual species to depth were similar for juvenile and adult fish. Number of species, CPUE and BPUE of all species except perch increased heading towards the tributary and peaked close to or at the tributary part of the reservoir. Responses of juvenile fish to distance from the dam differed from that of adult fish. Structure of fish community differed in benthic and pelagic habitats with species preferably occupying epipelagic (bleak, asp, rudd and juvenile bleak, roach and bream) or littoral waters (perch, pikeperch, ruffe, roach, bream and juvenile percids). This study showed that fish distribution in the reservoir followed distinct patterns, which were probably shaped by a combination of physiological constraints plus trade-off between food resources and competition.     

Habitat partitioning among three predatory fish in a temperate reservoir

The diversity of predatory species plays a key role in ecosystem functioning but our understanding of the mechanisms underlying their coexistence is limited, particularly in freshwater ecosystems. Northern pike Esox lucius, European perch Perca fluviatilis and pikeperch Sander lucioperca are three widespread predatory species in European lakes, where they often coexist. As potential competitors, we hypothesised that partitioning habitat is a determinant of species coexistence. This was tested by quantifying the variability of their habitat use in tracking adult individuals in the Bariousses reservoir (France, 86.6 ha, mean depth 7.1 m). Specifically, we investigated their distribution along the littoral–pelagic and depth axes along the daily cycle and across seasons. From littoral to pelagic waters were first found pike, then perch and finally pikeperch. Pike was the closest to the surface while pikeperch was the deepest. This general pattern was, however, variable across seasons with the three species located in the upper layer in summer during reservoir stratification. Individuals were more evenly distributed along the littoral–pelagic axis and closer to the bottom when water was mixing (autumn, winter). In summer, perch used more intensively in the pelagic zone during the daytime. Other species did not show any diel change of habitats. Our results highlighted that species coexistence is associated with habitat partitioning among these three predators, with perch showing a more variable behaviour regarding habitat characteristics. Now more than ever, in the context of global change which modifies habitats, it is of crucial importance to understand the coexistence mechanisms of species that shape ecosystems.     

A comparative study of three different methods for assessing fish communities in a small eutrophic lake

Sound decisions on the management of fish stocks depend on knowledge about the species composition, number, biomass and size structure of existing populations. Accordingly, the ability to make solid population estimates is essential. In this study, a 2.15 ha lake was completely drained and the total number of fish was recorded and amounted to 180,915 individuals divided into seven species having a total weight of 1,395 kg. Before the draining, three commonly used methods in fish surveys were applied: multi‐mesh gillnets, point abundance sampling by electrofishing (PASE) and mark–recapture. Following the determination of the actual number and size distribution of each species, we evaluated the efficiency of the methods and found that gillnets caught a relatively high number of species (five out of seven) and thus proved to be the best tool for mapping species richness. However, gillnets were size selective towards larger individuals of perch (Perca fluviatilis) and did not catch roach (Rutilus rutilus) <5 cm. In contrast to gillnets, PASE was very effective at catching YOY fish in the shore zone but selected for larger‐sized roach. In sum, gillnetting proved to be the most accurate method for estimating species composition, PASE also being useful. Overall, mark–recapture provided relatively good estimates of population size but small‐sized (<11 cm) roach proved not to be well suited for mark–recapture surveys. We conclude that the best method(s) surveying fish stocks depends on various factors such as target species, size distribution and the purpose of the survey.     

Impaired littoral energy pathways cause a shift to pelagic resources by fish in recovering lake food webs

Fish communities may increase in biomass and productivity due to energy subsidies from the littoral invertebrate community. In lakes recovering from acidification and metal contamination, such as those in Sudbury, Ontario, Canada, impaired benthic invertebrate communities (i.e., low diversity with higher abundance of small‐bodied taxa) allowed a critical test of the role of these littoral pathways on fish diet. We compared fish abundance, diversity, diet and biomass in eight recovering and eight reference lakes and related availability of the main littoral and pelagic invertebrate groups to fish diet regime using stable isotope analysis. A Bayesian mixing model (MixSIR) was used to estimate diet likelihood, and convex hull analysis was used to estimate trophic niche space of fish communities. Fish biomass did not differ between impaired and reference lakes despite substantial differences in potential diet. Fish depended strongly on littoral benthos in the reference lakes but consumed more pelagic food in the impaired lakes. The trophic niche of the focal, most common fish species (i.e., yellow perch, smallmouth bass, pumpkinseed and brown bullhead) was larger in the impaired lakes. We attributed these differences to low diversity at the highest trophic levels of fish communities in the impaired lakes as well as to depauperate benthic invertebrate communities. In contrast to the food webs of most temperate lakes, fish in impaired lakes preyed less on littoral invertebrates yet still managed to maintain a reference lake level of biomass standing crop by relying more on pelagic resources – macro zooplankton such as Chaoborus.     

Fish fauna in a Protected Greek lake: biodiversity, introduced fish species over a 80-year period and their impacts on the ecosystem

Abstract –  Lake Pamvotis is a tertiary natural shallow lake located in the NW of Greece. For biogeographical reasons, it historically contained only four fish species: two endemics ( Phoxinellus epiroticus , Squalius pamvoticus ), one native to the West Greece ( Barbus albanicus ) and the ubiquitous Anguilla anguilla . These species were almost exclusively present in the lake before 1920, forming remarkable commercially exploited populations. From the 1930s through the 1990s, 20 species were introduced into the lake for purposes of eutrophication control or to enhance the fishery. These introductions, some of them accidental have led to the apparent decline, even loss, of the native species. During the last three decades the fish fauna of the lake has shifted from the native, clear-water species to a predominance of introduced species, mainly those adapted for turbid eutrophic water ( Rutilus panosi , Cyprinus carpio and Carassius gibelio , and several Asian cyprinids). The current fish assemblage in the lake is dominated by introduced species particularly the Mosquitofish ( Gambusia affinis ) and the Lourogobios ( Economidichthys pygmaeus ) in the littoral zone, the benthopelagic allogynogenetic Prussian carp ( C. gibelio) , and the opportunistic Trichonis roach ( R. panosi ), in the pelagic zone.     

Whole-lake influences of littoral structural complexity and prey body morphology on fish predator–prey interactions

Abstract –  We used a longline tethering method to examine the relationship between prey refugia, prey body morphology and the location and magnitude of predation mortality within an individual lake and among three lakes that differed in coarse woody habitat (CWH) and aquatic macrophyte abundances. Predation events were lowest in the macrophyte and/or CWH refuges, peaked at or just beyond the refuge edge and declined in pelagic portions of the lakes. Predation risk at the refuge edge and just beyond was positively correlated with the abundance of littoral refuge. In contrast, predation events within the refuge and in the pelagic zone were negatively correlated with littoral refuge abundance. Deep-bodied and spiny prey morphologies were less vulnerable to predation than fusiform prey lacking fin spines. The structural complexity of littoral zones and prey fish body morphology may influence the outcome of predator–prey interactions and ultimately determine fish species assemblages in lakes.     

Temperature‐ and light‐dependent ratio of energy gain to metabolic costs explains spatial and temporal habitat use of zooplanktivorous fish

Understanding the forces that drive habitat selection of species in communities is important in both ecology and evolution. In nature, species face variation in competition, predation and physical characters among habitats. Vendace (Coregonus albula (L.)) is a specialised zooplanktivorous fish predominantly using deeper water in lakes during summer, while roach (Rutilus rutilus (L.)) uses mainly the shallow littoral zone as well as the upper layer of the pelagic zone. To understand mechanisms behind habitat use of these species, I first conducted a predation experiment to investigate their sensitivity to predation by perch (Perca fluviatilis L.). Second, I performed a foraging experiment using different temperature and light treatments. I then used metabolic calculations to estimate energetic costs when foraging. I found no difference between species regarding sensitivity to predation. Vendace was the most efficient forager on zooplankton but also swam faster spending more energy compared to roach. Roach had a comparatively high metabolic rate in the lowest temperature, where their foraging efficiency was lowest. The energy gain ratio at 6°C was highest for vendace, while it was lowest for roach. In the highest temperature (18°C) and the lowest light level (1 lux), both species were similar in their energy gain ratio. The relative energy gain ratio provides a mechanism to explain habitat distribution for the two species. An increased understanding of the role of metabolism in combination with biotic interactions and habitat use may help to foresee effects of environmental change for different species.     

Annual and seasonal dynamics of fish in the brackish-water Matsalu Bay, Estonia

Abstract –  Matsalu Bay in the Väinameri (West-Estonian Archipelago Sea) is a relatively shallow but large bay. It is the only real delta estuary in the northern Baltic Sea. The fish fauna is strongly dominated by freshwater species such as perch Perca fluviatilis , roach Rutilus rutilus , white bream Blicca bjoerkna and rudd Scardinius erythrophthalmus . The study presents CPUE data describing annual and seasonal variations in the species composition, abundance and spatial distribution, based on 1 year of monthly samplings from August 2001 to August 2002 and routine coastal monitoring (in late July–early August) between 1994 and 2003. The bay can be divided into three rather distinct parts. The fish fauna of the inner part (salinity 0–2 p.p.t.) is different from two open parts and resembles that of freshwater lakes. In the openmost part (salinity usually 4–6 p.p.t.) some marine species, such as herring Clupea harengus membras , flounder Platichthys flesus and eelpout Zoarces viviparus may seasonally be abundant. Species composition and abundance of most species varies seasonally and interannually. Only the abundance of few species such as pike Esox lucius and pikeperch Stizostedion lucioperca was not impacted by seasonality. The most dynamic period in the bay is spring, when several species (some of them very rare in summer, such as herring and smelt Osmerus eperlanus ) enter the bay for spawning.     

Can underwater refuges protect fish populations against cormorant predation? Evidence from a large‐scale multiple pond experiment

Artificial structures can protect fish against predation by cormorants (Phalacocorax spp.). However, their effectiveness in larger water bodies with different fish communities in the presence of natural vegetation still needs to be explored. Using a large‐scale field experiment with 24 ponds stocked with differently composed fish communities, the present study investigates the extent to which the effect of artificial refuges on fish is species‐specific and determined by the characteristics of the fish community. This study provides strong experimental evidence for artificial refuges protecting fish against predation from cormorants, even in the presence of submerged vegetation. The effect of refuges was, however, highly species‐specific and depended on the composition of the fish community. Strong positive effects of refuges on rudd, Scardinius erythrophthalmus (L.), and roach, Rutilus rutilus (L.), populations were observed, especially in ponds where these species dominated. Overall, the total biomass of young‐of‐the‐year, 1‐year‐old and adult rudd and roach was on average 500, 7 and 15 times lower in ponds without than in ponds with refuges, respectively. No effect of artificial refuges on other fish species was found. This study indicates that artificial refuges can facilitate the coexistence of predation vulnerable fish populations with cormorants in lakes and ponds.     

Assessing vulnerability of New Zealand lakes to loss of conservation value from invasive fish impacts

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Resource use of crucian carp along a lake productivity gradient is related to body size,predation risk,and resource competition

Generalist fish species can feed on a wide resource spectrum and across trophic levels depending on resource availability and trophic interactions. Crucian carp (Carassius carassius) represents a good candidate species to investigate variation in the trophic ecology of generalist fish as it can be found in highly variable fish communities and its resource use is well documented. In this study, we explored the trophic ecology of crucian carp at the individual and population levels using stable isotope and gut content analysis. We tested if trophic resource use varied according to lake productivity, predation risk, intra- and interspecific competition, or individual fish size. We found that crucian carp resource preference was highly variable among and within lakes. In predator-free lakes, small crucian carp occurred in high densities, showed increased interindividual specialisation, and relied mainly on pelagic zooplankton. In presence of predators, large crucian carp occurred in low densities and included greater proportions of benthic macroinvertebrates in their diet. This shift in resource use was further favoured in productive, shallow lakes where littoral prey were probably abundant. Resource partitioning was an important factor determining crucian carp niche use, as fish had higher trophic position in absence of other cyprinids. Crucian carp showed highly dynamic resource use and food preferences in response to variable environmental conditions. Overlooking complex diet preferences of generalist fish may lead to an oversimplification of freshwater community dynamics.     

Reproductive ecology of two cyprinid fishes in an oligotrophic lake near the southern limits of their distribution range

Abstract –  The reproductive activities of two dominant fish species, roach Rutilus rutilus and rudd Scardinius erythrophthalmus were investigated from August 2000 to July 2002 in an oligotrophic lake, Lake Sapanca, in the northwest of Turkey. A total of 907 roach and 593 rudd individuals were examined. Macroscopic appearance of ovary, gonadosomatic index and oocyte diameter frequency distribution suggested that the spawning season of roach and rudd were from early April (13 °C) to late May (18 °C) and from early May (17.5 °C) to late June (23.5 °C), respectively. The spawning cycles of both species showed the same tendency in two successive years. The estimated mean total lengths at first maturity ( L _T50) were 12.26 and 14.98 cm for male and female roach, respectively, and 7.12 and 8.10 cm for male and female rudd, respectively. The sizes at 50% maturity corresponded to 3 years for roach and 1 year for rudd, for both sexes. The ratio of males to females was 1.3:1 and 1.2:1 for roach and rudd, respectively. The average condition factor K was 1.21 for rudd and 1.14 for roach, and it was not affected by sex or reproductive effort. Absolute fecundity varied from 7829 to 95,387 eggs between ages 3 and 9 for roach and from 1807 to 35,629 eggs between ages 1 and 7 for rudd. Comparisons of these results with earlier studies from more northern regions revealed that spawning of both species in this area started earlier, but spawning temperature and the duration of spawning was approximately the same.     

Trophic interactions between introduced lake trout (Salvelinus namaycush) and native Arctic charr (S. alpinus) in a large Fennoscandian subarctic lake

Introduced fishes may have major impacts on community structure and ecosystem function due to competitive and predatory interactions with native species. For example, introduced lake trout (Salvelinus namaycush) has been shown to replace native salmonids and induce major trophic cascades in some North American lakes, but few studies have investigated trophic interactions between lake trout and closely related native Arctic charr (S. alpinus) outside the natural distribution of the former species. We used stomach content and stable isotope analyses to investigate trophic interactions between introduced lake trout and native Arctic charr in large subarctic Lake Inarijärvi in northern Finland. Both salmonids had predominantly piscivorous diets at >280 mm total length and were mainly caught from the deep profundal zone. However, lake trout had a more generalist diet and showed higher reliance on littoral prey fish than Arctic charr, whose diet consisted mainly of pelagic planktivorous coregonids. According to length at age and condition data, lake trout showed slightly faster growth but lower condition than Arctic charr. The results indicate that introduced lake trout may to some extent compete with and prey upon native Arctic charr, but currently have only a minor if any impact on native fishes and food web structure in Inarijärvi. Future monitoring is essential to observe potential changes in trophic interactions between lake trout and Arctic charr in Inarijärvi, as well as in other European lakes where the two salmonids currently coexist.     

Variation in larval fish communities: implications for management and sampling designs in reservoir systems

Understanding spatial and temporal patterns in larval fish distributions is important for investigating factors related to recruitment variability and for developing sampling methodologies. Variation in species diversity (Shannon index) and density (number 100 m^?3) of larval fishes from nearshore and pelagic habitats in Glen Elder Reservoir, KS, USA, was assessed during 1999 and 2000. Species diversity and densities of individual taxa were significantly higher (P < 0.05) in nearshore habitats than pelagic habitats during both years. Larval fish densities were generally higher in the year 1999 with higher spring water levels. Gizzard shad Dorosoma cepedianum (Lesueur) was the most abundant taxon in nearshore and pelagic habitats during both years. In addition, larval gizzard shad was sampled for the longest duration in the upper water column and overlapped with all taxa collected. High densities of gizzard shad in both habitats suggested that the potential for competitive interactions was high, especially given the lack of aquatic vegetation and habitat heterogeneity in littoral habitats. Most of the variation in larval fish diversity and density was because of temporal variation. If sampling effort must be limited and the purpose of sampling is to monitor larval diversity or density, attention should focus on sampling the same location over long time periods rather than across a variety of sampling locations.     

Environmental determinants of fish community structure in gravel pit lakes

Gravel pit lakes are increasingly common, and there is an urgent need to better understand the functioning of these artificial and disconnected ecosystems. However, our knowledge of the environmental determinants of fish community structure within these types of lakes remains poor. In this study, we quantified the taxonomic diversity, fish species and life‐stage composition in 17 gravel pit lakes sampled in 2012 and 2013 located in south‐west France to determine the potential role of environmental variables (i.e. lake morphology, productivity, water quality and human‐use intensity) as drivers of fish community structure and composition. Our results demonstrated that fish community structure significantly differed between gravel pit lakes, and we notably found that lakes managed for angling hosted higher levels of taxonomic diversity. We also found that young and large lakes supported higher native species biomass and were dominated by native European perch (Perca fluviatilis). Older, smaller and more productive lakes, located closer to the main urban area, supported a higher biomass of non‐native species such as largemouth bass (Micropterus salmoides). Native and non‐native sport fishing species such as northern pike (Esox lucius), pikeperch (Sander lucioperca), common carp (Cyprinus carpio) and cyprinid prey species were positively associated with fishery management effort, suggesting that management practices play also a critical role in shaping fish species composition. Overall, our study demonstrated that fish community composition followed a predictable shift along environmental gradients associated with the maturation of gravel pit lakes and the associated human practices.