Can artificial plant beds be used to enhance macroinvertebrate food resources for perch (<Emphasis Type="Italic">Perca fluviatilis</Emphasis> L.) during the initial phase of lake restoration by cyprinid removal?

Restoration of shallow lakes to a clear-water state, often characterized by high submerged macrophyte cover and a high proportion of piscivores such as perch, Perca fluviatilis L., frequently involves removal of a large proportion of the zoobenthivorous fish, such as bream, Abramis brama L., and roach, Rutilus rutilus L. (i.e. biomanipulation). However, establishment of submerged macrophytes is often delayed following fish removal. This is unfortunate because plant beds typically host high densities of the macroinvertebrates constituting the diet of small perch and thus help perch to go through the bottleneck from feeding on macroinvertebrates to feeding on fish. Establishment of artificial plant beds may be a useful tool to enhance macroinvertebrate population growth and thus food resources for small perch until the natural plants have established. To investigate this restoration option, we studied during two growing seasons (June–October) the composition and abundance of the macroinvertebrate community in artificial plant beds installed in shallow Lake Væng (Denmark) comprising the initial phase of a biomanipulation effort by fish removal. Lake areas with artificial plant beds exhibited substantially higher macroinvertebrate densities than the lake bottom. This suggests that artificial plant beds may be used as feeding grounds for small perch, similarly to the well-known refuge effect for zooplankton against fish predation. In this way, artificial plant beds could help maintain a clear-water state during the transient period when natural submerged vegetation is not yet established in the lake.     

Is coarse woody debris in lakes a refuge or a trap for benthic invertebrates exposed to fish predation?

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Collapse of the native ruffe (<Emphasis Type="Italic">Gymnocephalus cernua</Emphasis>) population in the Biesbosch lakes (the Netherlands) owing to round goby (<Emphasis Type="Italic">Neogobius melanostomus</Emphasis>) invasion

We investigated the change in benthic fish communities in three artificial lakes of the Biesbosch area in the Netherlands between two time periods: before and after the invasion of round goby (Neogobius melanostomus). Native ruffe (Gymnocephalus cernua), the dominant species in benthic gillnet and littoral beach seining catches before the invasion, almost completely disappeared in all lakes only 2 years after the invasion. We found a significant increase in 0 + perch (Perca fluviatilis) and, in some lakes, pikeperch (Sander lucioperca) abundance in gillnet catches after invasion. In the post-invasion period, the 0 + fish community was dominated by perch, and the older fish community was dominated by round goby. The species richness of 0 + fish increased in the post-invasion period owing to the invasion of gobiids. However, it did not change for older fish between periods. Our results clearly show that, owing to a similar benthic lifestyle and high niche overlap, ruffe was the only species negatively influenced by the round goby invasion. The competitive superiority of round goby over ruffe is so strong that the once-dominant species of the overall benthic fish community collapsed after only a few years of coexistence.     

Patch based predation in a southern Appalachian stream

Streams are characterized by high degrees of patchiness that could influence the role of predators in these systems. Here we assess the impact of predatory benthic fishes on benthic macroinvertebrate density, biomass, and community structure at the patch scale in a fourth order stream in the southern Appalachians. We tested the role of predation in two different patch types: patches inhabited by adult mottled sculpin ( Cottus bairdi ) and random patches. We placed 30 basket pairs (one open to fish predation, and one from which fish predators were excluded) in the streambed at each patch type. We also tested for potential basket effects by setting up a basket control area. Although there was some evidence of basket artifacts on macroinvertebrate density in sculpin patches, these artifacts were not consistent and we do not feel that they affected our results because predators did not affect macroinvertebrate density. In random patches, predation did not significantly affect macroinvertebrate density or biomass. Predators significantly reduced macroinvertebrate biomass in sculpin patches but did not affect prey density. When the data-set was size-limited to exclude macroinvertebrates too large for consumption by sculpin, macroinvertebrate biomass did not differ significantly between exclusion and open baskets. This suggests that sculpin can reduce macroinvertebrate biomass through a combination of consumption and by predator-induced emigration of large macroinvertebrates into areas that are protected from sculpin. In addition, invertebrate predator biomass was higher in predator exclusion baskets in sculpin patches indicating that predation pressure remained high in the exclusion baskets despite fish exclusion. These results illustrate the heterogeneity of streams and the effect of small-scale differences (e.g. location of predators' territories) on local processes. Experiments that utilize these differences can provide insights into these stream processes.     

Climate and productivity shape fish and invertebrate community structure in subarctic lakes

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Associations Between Macroinvertebrates and <Emphasis Type="Italic">Paralemanea mexicana</Emphasis>, an Endemic Freshwater Red Alga from a Mountain River in Central Mexico

Macrophytes are common inhabitants of lotic environments and, depending on their morphological traits, possess adaptations that provide shelter to aquatic invertebrates against strong river flow and predators. They may also be used as a food source by macroinvertebrates. The main goal of this study was to determine the relationship between the red alga Paralemanea mexicana and its role as a shelter and/or food source for lotic macroinvertebrates. We also conducted research on the role of microhabitat and morphological variations of the alga in determining macroinvertebrate taxon abundance, diversity, and functional group composition in a high-current velocity river. Results showed that changes in cover and morphology of P. mexicana were mostly correlated with river current velocity, irradiance, and seasonal variation. In turn, these were related to changes in abundance and diversity of the associated macroinvertebrate community. In addition, six macroinvertebrate functional feeding groups were evaluated for associations with the red alga: filtering and gathering collectors, piercers, scrapers, herbivore shredders, and predators. The results showed that the Trichoptera Hydroptilidae genera Ochrotrichia and Metrichia use P. mexicana as a food source and case-building material. The Trichoptera Glossosomatidae Mortoniella uses the alga as a substrate. The biotic interactions between P. mexicana and associated macroinvertebrates reveal the importance of macrophytes as purveyors of substrate, as food and shelter for macroinvertebrates, and also as promoters of macroinvertebrate community diversity. In addition, it was shown that macroinvertebrate herbivory likely facilitates vegetative propagation of the red alga through increased release and germination of carpospores and new gametophytes.     

Scale-dependant control of motile epifaunal community structure along a coral reef fishing gradient

Large-scale fishing is mostly conducted using towed gears that reduce the biomass and diversity of benthic invertebrates. However, it is impossible to differentiate between the physical disturbance effect of towed gears from the effect of fish predator removal upon benthic invertebrate communities. Here we explore the impact of fish removal alone on the community structure of small motile coral reef invertebrates (epifauna) along a subsistence fishing intensity gradient in the Lau group, Fiji. We deployed settlement plates at three areas in each of six fishing grounds and examined the density and class richness of the motile epifaunal communities and the associated algal communities in relation to the structure of fish and benthic communities. Motile epifaunal density was unrelated to fishing intensity. However, at smaller inter-area scale (0.5-10 km) motile epifaunal density was negatively related to plate algal biomass, whereas at the larger inter-fishing-ground scale (4-180 km) motile epifaunal density was positively related to the rugosity (substrate complexity) of the surrounding benthos. The class richness and diversity (Margalef's d) of motile epifaunal communities were negatively related to fishing intensity, but unrelated to grazing intensity, rugosity or algal biomass at either scale. Benthic community structure varied significantly with fishing intensity; hard-coral cover was lower and turf-algal cover was higher at high fishing pressure. The variation in benthic community structure was associated with variation in fish community structure, which in turn varied with fishing intensity. Motile epifaunal community structure upon plates was linked to the structure of the surrounding benthic community, but was not directly linked to the plate algal community. We suggest the decline in richness of the motile epifauna community along the fishing gradient is attributable to either to exploiter-mediated coexistence or the reduction in ‘habitat quality’ of the surrounding benthos. At the large spatial scale substrate complexity is the key determinant of motile epifaunal density, suggesting predation by fishes plays an important structuring role at this scale. Assuming that rugosity is inversely related to predation risk then this study represents the first evidence for spatial-dependence on the top-down (predation) vs. bottom-up (algal biomass) control of community structure. We argue fisheries exploitation, in the absence of a physical disturbance can negatively influence motile epifaunal community structure at large spatial scales.     

Stream inflow and predation risk affect littoral habitat selection by benthic fish

1. We examined small, fishless headwater streams to determine whether transport of macroinvertebrates into the littoral zone of an oligotrophic lake augmented food availability for Cottus asper, an abundant predatory fish in our study system. We sampled fish and macroinvertebrates during the recruitment and growth season of 2 years, either monthly (2004) or bi‐monthly (2005), to observe whether stream inputs increased prey availability and whether variation in total macroinvertebrate biomass was tracked by fish. 2. Observations from eight headwater streams indicated that streams did not increase the total macroinvertebrate biomass in the shallow littoral zone at stream inflows, relative to adjacent plots without stream inputs (controls). The taxonomic composition of stream macroinvertebrates drifting toward the lake differed from that in the littoral lake benthos itself, although there was no evidence of any species change in the composition of the littoral benthos brought about by stream inputs. 3. Although streams made no measurable contribution to the biomass or taxonomic composition of the littoral macroinvertebrate benthos, there was substantial temporal variation in biomass among the eight sites for each of the (n = 7) sample periods during which observations were made. Variation in total biomass was primarily a function of bottom slope and benthic substrata in the lake habitats. Dominant taxonomic groups were Baetidae, Ephemerellidae (two genera), Leptophlebiidae, Chironomidae (three subfamilies) and Perlodidae, although we did not determine the specific substratum affinities of each taxon. 4. Mixed effects linear models identified a significant interaction between macroinvertebrate biomass and plot type (stream inflow vs. control) associated with fish abundance. Across the observed range of macroinvertebrate biomass, fish showed a significant preference for stream inflows, but more closely tracked food availability in the controls. For young‐of‐the‐year (YOY), a negative effect of temperature was also included in the model, and we observed lower temperatures at stream inflows. However, abundance of predatory adults affected habitat selection for YOY. Lake‐bottom slope also accounted for variation in abundance in both fish models. 5. Our results suggest that the effect of fishless headwater streams on downstream fish may not always be through direct delivery of food. In this study system, fish preferred stream inflow plots, but this preference interacted with macroinvertebrate biomass in a manner that was difficult to explain. For YOY, predation risk was related to the preference for stream inflows, although the specific factor that mitigates predation risk remains poorly understood.     

Growth,morphological variation and ontogenetic niche shifts in perch (<Emphasis Type="Italic">Perca fluviatilis</Emphasis>) in relation to resource availability

Despite the common occurrence of ontogenetic niche shifts, their consequences for morphological adaptations have been little studied. To address this question, we studied morphological adaptations related to ontogenetic niche shifts in Eurasian perch (Perca fluviatilis) in eight lakes that varied in density of benthic resources and planktivorous fish biomass. Perch start to feed on pelagic zooplankton, then shift to benthic resources at intermediate sizes, and finally, when large enough, mainly feed on fish. These three functional niches over ontogeny are expected to set constraints on the morphology and size-specific growth of perch. The growth of perch was negatively related to planktivorous fish biomass in the zooplanktivorous niche, but positively related to planktivorous fish biomass in the piscivorous niche. The number of gill rakers of perch was negatively related to the biomass of planktivorous fish, providing evidence for the occurrence of character displacement as a consequence of competition in the zooplanktivorous niche. Perch in lakes with low densities of predator-sensitive macroinvertebrates had greater body height measurements and a larger mouth early during ontogeny. This pattern is suggested to be a result of a selection for increased efficiency in the benthic niche when the availability of benthic resources is low. Perch in lakes with a high biomass of planktivorous fishes had fusiform body morphology, a thicker tail and a larger mouth then the average piscivorous perch. The different responses of perch morphology to variation in the availability of benthic resources compared to variation in planktivore biomass are suggested to be partly because the availability of the former resource to a larger extent is set by abiotic conditions (humic content). We suggest that the key factors affecting size-specific growth and body morphology of perch in the system studied are the availability of resources in the benthivorous and piscivorous niches. We also provide evidence for morphological trade-offs, especially between the benthivorous and the piscivorous ontogenetic niches. Received: 6 July 1999 / Accepted: 8 September 1999     

The impact of predation by burbot (<Emphasis Type="Italic">Lota lota</Emphasis> L.) on the macroinvertebrate community in the littoral zone of a large lake

We studied the effects of predation by juvenile burbot (Lota lota) on the macroinvertebrate community in mesocosm experiments in the stony littoral zone of Lake Constance, a large prealpine lake in Central Europe. Although the growth data of the burbot suggest that the benthivorous fish exerts strong predation pressure on the invertebrate community, the predicted level of consumption is only poorly reflected by changes in biomass and abundance of most invertebrate prey taxa. High exchange rates of the prey between the mesocosms and the ambient littoral environment apparently masked the true predation effects of fish. Also, life-history events such as hatching or synchronised emergence of larvae led to temporal effects that obscured the impact of predation. However, for the dominant prey organism, the freshwater amphipod Gammarus roeseli, direct lethal effects appeared when its migration was limited. When exchange with the ambient littoral zone was possible, abundance and biomass of G. roeseli were unexpectedly high in the mesocosms stocked with burbot, indicating behavioural responses to fish presence. During the experiment, the burbot gained ca. 15% in length and about 60% in body mass. According to stomach content analyses at the end of the experiment, the ingested prey consisted mainly of relatively large and abundant invertebrates. Our study indicates that predation by juvenile burbot should be an important factor in structuring the benthic invertebrate community in terms of qualitative and quantitative effects.     

The impact of the invasive Ponto-Caspian amphipod <Emphasis Type="Italic">Pontogammarus robustoides</Emphasis> on littoral communities in Lithuanian lakes

The intentionally introduced Pontogammarus robustoides is the most successful amphipod invader of Lithuanian inland waters and has become established in large lakes. Its impact on littoral invertebrate communities was studied by comparing similar habitats across lakes that harbour or are devoid of the invader. In habitats where P. robustoides is well established and numerous, it significantly reduces species richness and community diversity. Moderate pontogammarid density in habitats that can sustain the native gammarid Gammarus lacustris, however, revealed no negative impact on diversity metrics. Among the lakes studied, the benthic biomass did not differ in invaded and uninvaded habitats. The biomass of indigenous invertebrates (excluding chironomids, which exhibited high lake-specific biomass variation) was lower in the places with well-established P. robustoides. A detrimental impact was observed upon the native isopod Asellus aquaticus and a negative correlation with most of the higher taxa of native invertebrates. In the invaded lake habitats that favour P. robustoides, a change in community structure and a decrease in diversity up to twofold or more are to be expected.     

Ethoxyresorufin-O-deethylase activity and content of polycyclic aromatic hydrocarbon metabolites in bream <Emphasis Type="Italic">Abramis brama</Emphasis> L. and perch <Emphasis Type="Italic">Perca fluviatilis</Emphasis> L. from the Sutka River (Rybinsk Reservoir)

This article presents the results of a comparative analysis of hepatic ethoxyresorufin-O-deethylase (EROD) activity and content of biliary polycyclic aromatic hydrocarbon (PAH) metabolities in bream Abramis brama L. and perch Perca fluviatilis L. from a region with background levels of organic xenobiotics. Statistically significant interspecific differences are revealed. The results of the study should be considered in studies on the effects of contaminants in fish.     

Influences of fish on food web structure and function in mountain lakes

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Effects of temperature,pH, and heavy metals (copper,zinc) upon proteinase activities in digestive tract mucosa of typical and facultative piscivorous fish

Species-specific differences and effects of natural (temperature, pH) and anthropogenic (zinc, copper) factors upon activities of casein-and hemoglobinlytic proteinases have been studied in the gut mucosa in pike (Esox lucius), zander (Stizostedion lucioperca), burbot (Lota lota) and perch (Perca fluviatilis). It is revealed that the intestine mucosa proteinase activities differ considerably from those ones in the fish stomach mucosa and that studied factors affect these activities to different extents. The stomach mucosa proteinases are most tolerant to the effects of studied factors, which is especially true for pike. The least tolerant are the intestine caseinlytic proteinases (except for the enzymes in perch). The effects of temperature, pH, and heavy metals depend on fish species, factors’ combinations (mono-factor, bi-factor, or multi-factor action) and on factor variability.     

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.     

Effects of an invasive crayfish on the littoral macroinvertebrates of large boreal lakes are habitat specific

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Population traits and colonization success of non-native Eurasian perch (<Emphasis Type="Italic">Perca fluviatilis</Emphasis>) 35 years after its first appearance in the Mediterranean Lake Skadar

Abundance, spatial distribution, population structure and growth of non-native Eurasian perch (Perca fluviatilis) were examined in the Lake Skadar, a large, unstratified lake in the Mediterranean region inhabited by several endemic, rare and threatened fish species. Fish were caught in the infra-littoral and littoral habitats of the northern and central parts of the lake using multi-mesh gill nets over three consecutive years. Eurasian perch was among the most dominant species in both habitats. The mean relative abundance is - NPUE (0.556 individuals m^?2) and relative biomass is - WPUE (23.56 g m^?2) (years and habitats combined). Its population consisted primarily of individuals in first, second and third year of life span (0+???2+ age classes). The youngest fish in first and second year showed low mortality rates, older fish in third, fourth, fifth and sixth year of life span (age classes 2+???5+) experienced great losses, with mortality rates ranging from 0.6–1.0. The von Bertalanffy growth curve formula, expressing the expected total length L _t ?=?36.98 (1-e^{-0.23 (t?+?1.018)}) with growth performance value Φ ^’ ?=?2.50, revealed fast growth-in-length. Length-to-weight relationship was W_t?=?6.2?×?10^?3 L _t ^3.27, indicating positive growth-in-weight. It has been concluded that 35 years after the first finding in the Lake Skadar, Eurasian perch adapted well to local conditions in this Mediterranean environment, with the life span changed in favor of fast growth and early maturation.     

Response of benthic macroinvertebrates to whole-lake, non-native fish treatments in mid-elevation lakes of the Trinity Alps, California

Introduced fish reduce the abundance and diversity of native aquatic fauna, but the effect can be reduced in complex habitats. We manipulated fish populations in forested mountain lakes to determine whether or not fish affected benthic macroinvertebrate composition across lakes with differing habitat complexity. We compared abundance, biomass, body-length, and community structure of benthic macroinvertebrates from 16 lakes with three treatments (fish stocked, suspended stocking, fish removed) and unstocked fishless “controls”. Over 4 years, we assessed the relative importance of fish and environmental variables influencing the composition of benthic macroinvertebrates. Control lakes had the greatest overall abundance of macroinvertebrates when chironomid midges were excluded. Abundances of insects in the clinger/swimmer functional group and caddisflies were greatest in the control lakes but were primarily influenced by habitat variables including the availability of aquatic vegetation and wood. Total biomass and mean body length of macroinvertebrates were not affected by treatment. Taxon richness of macroinvertebrates was about 40% greater in the control lakes compared to the treatment lakes but did not differ among treatments. Our results suggest that fish reduce susceptible macroinvertebrate richness and abundances, but that changes associated with alterations of fish composition are confounded by other factors in complex lake habitats.     

Weedbeds and big bugs: the importance of scale in detecting the influence of nutrients and predation on macroinvertebrates in plant‐dominated shallow lakes

1. The scale of investigations influences the interpretation of results. Here, we investigate the influence of fish and nutrients on biotic communities in shallow lakes, using studies at two different scales: (i) within‐lake experimental manipulation and (ii) comparative, among‐lake relationships. 2. At both scales, fish predation had an overriding influence on macroinvertebrates; fish reduced macroinvertebrate biomass and altered community composition. Prey selection appeared to be size based. Fish influenced zooplankton abundance and light penetration through the water column also, but there was no indication that fish caused increased resuspension of sediment. 3. There were effects of nutrients at both scales, but these effects differed with the scale of the investigation. Nutrients increased phytoplankton and periphyton at the within‐lake scale, and were associated with increased periphyton at the among‐lake scale. No significant effect of nutrients on macroinvertebrates was observed at the within‐lake scale. However, at the among‐lake scale, nutrients positively influenced the biomass and density of macroinvertebrates, and ameliorated the effect of fish on macroinvertebrates. 4. Increased prey availability at higher nutrient concentrations would be expected to cause changes in the fish community. However, at the among‐lake scale, differences were not apparent in fish biomass among lakes with different nutrient conditions, suggesting that stochastic events influence the fish community in these small and relatively isolated shallow lakes. 5. The intensity of predation by fish significantly influences macroinvertebrate community structure of shallow lakes, but nutrients also play a role. The scale of investigation influences the ability to detect the influence of nutrients on the different components of shallow lake communities, particularly for longer lived organisms such as macroinvertebrates, where the response takes longer to manifest.