Stable isotopes indicate that zebra mussels (Dreissena polymorpha) increase dependence of lake food webs on littoral energy sources

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Uncoupling strong predator–prey interactions in streams: the role of marginal macrophytes

Predation refugia can facilitate the coexistence of predators and prey within an ecosystem by weakening trophic links between the two. The marginal macrophytes of shallow lakes are used facultatively by zooplankton to escape fish predation, leading to the stabilisation of lentic food webs. Little is known about such a role for the marginal macrophytes of lotic systems. In this paper, we examine whether the marginal macrophytes of chalk streams help buffer the interaction between the freshwater shrimp, Gammarus pulex and a benthic fish predator, Cottus gobio, both characteristically abundant in such systems. Quantitative surveys were taken of Gammarus and bullheads in winter and summer in a southern English chalk stream. These indicated that Gammarus seasonally switched their habitat preference, from marginal macrophytes in summer, to midchannel habitats in winter. Bullheads exhibited an opposite trend, preferring midchannel habitats in summer and all habitats, particularly marginal macrophytes, in winter. Large Gammarus and precopula pairs were found almost exclusively in the margins in summer. This spatial separation between Gammarus and bullheads in summer, though not in winter, was reflected in bullhead diets, as determined by gut analysis. In field manipulation experiments, bullhead presence was the strongest factor explaining the between‐habitat distribution of Gammarus in both summer and winter, indicating that the habitat shift of Gammarus was driven by the distribution of fish. Other abundant invertebrate taxa, including Asellus, mayflies and chironomids, exhibited little avoidance of habitat patches of high bullhead density. We conclude that marginal macrophytes in chalk streams can potentially facilitate the coexistence of high densities of both Gammarus and bullheads by spatially separating predator and prey in summer. They may further allow large populations of Gammarus to persist in the presence of high bullhead density at stretch‐wide spatial scales, by reducing the predation by bullheads of large breeding and newly born individuals of Gammarus in summer.     

Temporal and spatial macrofaunal community changes along a salinity gradient in seagrass meadows of Akkeshi-ko estuary and Akkeshi Bay,northern Japan

Changes in water clarity (secchi disc transparency) in relation to the presence/absence of introduced, exotic fish, including rudd (Scardinius erythrophthalmus), tench (Tinca tinca), perch (Perca fluviatilis), brown bullhead catfish (Ameiurus nebulosus), goldfish (Carassius auratus), and koi carp (Cyprinus carpio) were determined for 49 small, North Island, New Zealand lakes. There was a negative association between water clarity and the presence of exotic fish independent of lake depth. Moreover, a ‘before-and-after’ comparison and examination of case-studies indicated that introductions of exotic fish reduce water clarity. The number of species introduced affected the relationship between lake depth and water clarity but the specific role of each species could not be distinguished because most of the lakes (83%) contained more than one exotic fish species. A model incorporating the known mechanisms by which planktivorous, benthivorous and herbivorous fish can influence water clarity in lakes showed that control over just one species or feeding guild may not result in an improvement in water clarity because of the additive and synergistic effects of different species on lake trophic processes. Handling editor J. Cambray     

Habitats and Littoral Zone Fish Community Structure of Two Natural Lakes in Southeast Brazil

The fish community and habitat structure in the littoral zone of lakes Pedra and Hortência, middle Rio Doce Valley, Brazil, were investigated in three sampling periods from August 1992 to May 1993. A total of 9106 fishes were collected, including 11 species in Lake Hortência and 12 species in Lake Pedra. Diversity was higher in Lake Hortência, although total fish abundance was the same for both lakes, with no significant temporal differences. The length distribution of the majority of fish species was similar between lakes. Geophagus brasiliensis was the dominant species (number/biomass) in all habitats of the two lakes in all three sampling periods. A striking difference in total fish abundance in relation to habitat types, which we attribute to physical variables, especially the abundance of macrophytes was observed. Negative impacts attributable to the local Eucalyptus sp. plantations were not detected in the fish community structure of Lake Hortência, which is probably due to the maintenance of the riparian vegetation contributing to similar physical environmental conditions in the littoral zone and high similarity in fish community structure between the two lakes. However, further studies on fish community structure and potential Eucalyptus impacts are suggested as necessary to improve conservation measures of the lakes and their fish faunas.     

Lake responses to long‐term disturbances and management practices

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A water level drawdown index for aquatic macrophytes in Nordic lakes

Many northern lakes are regulated to enhance hydropower production and flood protection. This bears hydromorphological pressures which are important factors causing lowered ecological status. Water level fluctuation triggers erosion on the shoreline and, depending on fluctuation range, also affects species composition or disappearance of sensitive aquatic macrophytes. We developed a water level-drawdown index (WI_c) for Nordic lakes using macrophyte data from 73 lakes with varying water level fluctuation in Finland, Norway and Sweden. The index is based on the ratio between sensitive and tolerant macrophyte species. The sensitive and tolerant species are identified based on a percentile approach, analysing the presence or absence of species along the winter drawdown range. The index correlates well with winter drawdown in Finnish and Norwegian lakes with strongest correlations with winter drawdown in storage lakes (lakes regulated for hydroelectric power and with a considerable winter drawdown). The WI_c-index is applicable in low alkalinity, oligotrophic and ice-covered lakes, and is suggested to be a useful tool to identify and designate heavily modified water bodies in Nordic lakes according to the European Water Framework Directive.     

Associations between water chemistry and fish community composition: a comparison between isolated and connected lakes in northern Sweden

1. The correlation between water chemistry, physical variables and fish community composition was examined in 40 small (≤30 ha) coastal lakes in northern Sweden. Twenty of the 40 lakes were isolated from other water bodies and 20 were connected to the Baltic Sea. Lakes were fished in summer, using three different methods. Water chemistry was sampled in late winter prior to ice‐out and pH was measured additionally in summer. 2. Our central question was whether water chemistry plays a greater role in the composition of fish communities in isolated lakes than in connected lakes, as isolated lakes cannot be recolonised once a species has become extinct. 3. Results indicate that winter anoxia affects community composition only in isolated lakes, whereas acidity is of importance in both connected and isolated lakes. Methane (indicating anoxia), was significantly correlated with variation in fish community composition in isolated lakes, and a group of variables that indicate anoxia (CH₄, pCO₂, inorganic carbon and dissolved oxygen) explained 24–34% of the variation. pH alone explained 12% of the variation in community composition for connected lakes and a group of variables indicating acidity (summer and winter pH and ANC) explained 10–20% of the variation in isolated lakes. Lake area was the most important physical variable, being significantly correlated with the variation in fish community composition in connected lakes. 4. In isolated lakes, the presence of pike (Esox lucius), perch (Perca fluviatilis) and roach (Rutilus rutilus) was associated with low CH₄. The occurrence of crucian carp (Carassius carassius) and roach was positively correlated with pH, and the crucian carp was apparently also affected by predation by pike and perch. In connected lakes the effect of anoxia was low, probably due to the possibility of recolonisation and pockets of oxygenated water, allowing pike and perch to persist and thereby limiting the distribution of crucian carp.     

Do introduced crayfish affect benthic fish in stony littoral habitats of large boreal lakes?

Invasive crayfish are spreading rapidly across Europe, where they are replacing the native crayfish species and impacting negatively on some other biota. Freshwater crayfish and many benthic fishes share similar habitat and food requirements and hence potentially compete for resources. In this study, we investigated impacts of the introduced signal crayfish (Pacifastacus leniusculus) on fish in stony littoral habitats of two large boreal lakes. We compared the littoral fish community composition and the densities of two common benthic fish species between sites with and without crayfish. To evaluate whether signal crayfish share the same food resources as benthic littoral fish or change their feeding habits, we used mixing models and trophic niche estimates based on analyses of stable isotopes of carbon and nitrogen. Both the community composition of littoral fish and the densities of benthic fish species were similar at sites with and without signal crayfish. Even though stable isotope signatures indicated strong dietary overlap between crayfish and benthic fish, the use of food sources and trophic niche widths of fish were not noticeably different between crayfish sites and non-crayfish sites. Our results suggest that, at current densities, the non-native signal crayfish does not have significant impacts on benthic fish in the stony littoral habitats of large boreal lakes.     

Effect of landscape factors on fish distribution in arctic Alaskan lakes

1. The distribution of species is affected by many factors operating at a variety of temporal and spatial scales in a heterogeneous landscape. In lakes, fish communities are dynamic, influenced by landscape‐level factors that control colonisation and extinction. 2. We used classification and regression tree (CART) analyses to quantify the importance of landscape‐level factors in determining the distribution of fish species in 168 arctic Alaskan lakes. Factors including lake size, depth, outflow gradient, distance to other lakes, lake order, altitude, river drainage and age of glacial surface were analysed. These factors could affect either access of fish to a lake (colonisation variables), or their survival in a lake that already had been colonised (extinction variables). 3. The presence of a species was predicted accurately in 78.4% ± 10.5% (mean ± SD) of cases, and absence in 75.0% ± 6.1% of cases. The relative importance of extinction versus colonisation variables varied with species. Extinction variables were most important for lake trout (Salvelinus namaycush) and slimy sculpin (Cottus cognatus), a mixture of extinction and colonisation variables was important for arctic char (Salvelinus alpinus), and colonisation variables were most important for arctic grayling (Thymallus arcticus) and round whitefish (Prosopium cylindraceum). 4. Ecological differences among species account for much of the difference in relative importance of colonisation versus extinction variables. In addition, stream piracy events have occurred over geologic time scales, which have resulted in lakes that are currently inaccessible but support relict fish populations. 5. Climate warming, currently occurring in the arctic, is likely to alter further the stream network, which could have dramatic effects on fish distributions by affecting access to isolated lakes or isolating lakes that are currently accessible.     

Lipid‐rich zooplankton subsidise the winter diet of benthivorous Arctic charr (Salvelinus alpinus) in a subarctic lake

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Littoral zone fish communities of two Florida lakes and a comparison with Michigan lakes

Synopsis The abundance and habitat distribution of littoral zone fishes in two small southern Florida lakes were quantified by underwater censuses. The bluegill (Lepomis macrochirus) and large-mouth bass (Micropterus salmoides) accounted for 75–80% of community biomass in both lakes; important coexisting species were predominantly benthic feeders in Lake Sirena and planktivores in Lake Annie. All species were largely confined to areas of macrovegetation which extended to a depth of 4 m in Lake Annie but only to 1.2 m in Lake Sirena. The differences in community structure were related to differences in habitat and also variation in water levels and benthic production.A comparison of community composition with that of small Michigan lakes indicated that similar numbers of species coexist in the littoral zones, despite a three-fold difference in the size of regional species pools. The majority of families and about 50% of the species were common to lakes in both regions; in addition a number of ecological analogues were noted. A major difference was that the small inshore species were members of the Cyprinodontiformes in Florida and Cyprinidae in Michigan. Together the largemouth bass and bluegill comprise similar community proportions in the two regions as do other major feeding groups. Lakes in the two regions that are similar in amount and distribution of vegetation exhibit greater similarity in fish communities than those within regions that differ in littoral vegetation.     

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

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Long-term changes in littoral fish community structure and resilience of total catch to re-oligotrophication in a large,peri-alpine European lake

1. The littoral zone of lakes is used as spawning, shelter, or feeding habitat for many fish species and hence is of key importance for overall lake functioning. Despite this, hardly any studies exist examining the long-term dynamics and response of the littoral fish community, composed mostly of juvenile fish, to environmental change. Here, we study the response of total catch per unit effort (CPUE) and individual species CPUE of such a community to 17 years of oligotrophication and examine whether the species responses can be characterised as synchronous or asynchronous.
2. We analyse a data set of beach seine catches carried out during morning and twilight, late spring and late summer at three sites in large and deep Lake Constance from 1997 to 2014. Generalised additive mixed models were used to explore changes in CPUE of the overall community and of the most frequently occurring species, and Kendall's W test was applied to examine whether the dynamics of fish species were synchronous or asynchronous.
3. Species-specific and total CPUE strongly differed between morning and twilight and between spring and summer indicating an important role of behavioural and life cycle adaptations of species for CPUE. In addition, also the CPUE of some species seeking shelter behind larger stones was lower at sites without these.
4. Total CPUE did not decline suggesting the overall abundance of littoral fish was resilient to declining nutrients. In contrast, fish community composition changed strongly during the study period due to increases in some species (dace, loach, perch) and decreases in others (bream, burbot, chub, ruffe), indicating response diversity of fish to oligotrophication. The type of community dynamics was scale-dependent, whereby significantly synchronous dynamics according to Kendall's W were observed when taking seasonal variability into account. In contrast, significantly asynchronous species dynamics were observed when only the low-frequency variability of species dynamics was considered separately for spring and summer time series.
5. Resilience of littoral fish total CPUE to oligotrophication might have important consequences for ecosystem dynamics and ecosystem services beyond the littoral zone. As small fish often impose strong predation pressure on zooplankton, their resilience might sustain a high top-down control on zooplankton resulting in a further reduction of zooplankton biomass. This could contribute to delayed food web responses and reduced growth of fish with oligotrophication.

     

Relationships between fish feeding guild and trophic structure in English lowland shallow lakes subject to anthropogenic influence: implications for lake restoration

The shallow lakes of Eastern England have been subject to intense anthropogenic pressures including nutrient enrichment and fish stocking. We sought to determine the relationships between fish community structure and other ecosystem characteristics in 28 of these lakes through classification of fish species into piscivorous, zooplanktivorous and benthivorous feeding guilds according to the literature. Canonical correspondence analysis produced clear associations between fish and ecosystem characteristics that generally agreed with other theoretical (e.g. the alternative stable states hypothesis) and empirical studies, but with some important differences. There was a striking lack of relationships between nutrients and other variables, indicating the importance of top-down rather than bottom-up processes as a structuring force in the generally eutrophic study lakes. The presence of submerged (and shoreline) vegetation was associated with a diverse assemblage of apparently co-existing piscivorous (principally pike Esox lucius) and zooplanktivorous species. Perch Perca fluviatilis, a significant predator in other studies, was unimportant and argued to be limited by water quality in the extremely shallow lakes. In contrast, the benthivorous fish guild (principally carp Cyprinus carpio, bream Abramis brama and tench Tinca tinca) essentially represented the inverse of the potential pelagic associations between piscivores/zooplanktivores and vegetation. The introduction of large benthivores to many study lakes could have precipitated a loss of submerged vegetation through direct uprooting during foraging, with the effect of simplifying the fish community being most acute where littoral vegetation was limited by other anthropogenic factors. It is implied that attempts to promote or restore submerged vegetation in these lakes would best target benthivorous species.     

Impacts of zebra mussels (Dreissena polymorpha) on isotopic niche size and niche overlap among fish species in a mesotrophic lake

Zebra mussels (Dreissena polymorpha) filter feed phytoplankton and reduce available pelagic energy, potentially driving fish to use littoral energy sources in lakes. However, changes in food webs and energy flow in complex fish communities after zebra mussel establishment are poorly known. We assessed impacts of zebra mussels on fish littoral carbon use, trophic position, isotopic niche size, and isotopic niche overlap among individual fish species using δ¹³C and δ¹⁵N data collected before (2014) and after (2019) zebra mussel establishment in Lake Ida, MN. Isotope data were collected from 11 fish species, and from zooplankton and littoral invertebrates to estimate baseline isotope values. Mixing models were used to convert fish δ¹³C and δ¹⁵N into estimates of littoral carbon and trophic position, respectively. We tested whether trophic position, littoral carbon use, isotopic niche size, and isotopic niche overlap changed from 2014 to 2019 for each fish species. We found few effects on fish trophic position, but 10 out of 11 fish species increased littoral carbon use after zebra mussel establishment, with mean littoral carbon increasing from 43% before to 67% after establishment. Average isotopic niche size of individual species increased significantly (2.1-fold) post zebra mussels, and pairwise-niche overlap between species increased significantly (1.2-fold). These results indicate zebra mussels increase littoral energy dependence in the fish community, resulting in larger individual isotopic niches and increased isotopic niche overlap. These effects may increase interspecific competition among fish species and could ultimately result in reduced abundance of species less able to utilize littoral energy sources.

     

Patterns of lake occupancy by fish indicate different adaptations to life in a harsh Arctic environment

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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.     

Annual winter water level drawdowns limit shallow‐water mussel densities in small lakes

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Assessing lake typologies and indicator fish species for Italian natural lakes using past fish richness and assemblages

In order to establish a fish-based typology of Italian lakes and identify possible reference and indicator fish species for each lake type, we analysed historical data on fish assemblages of all Italian natural lakes >0.5 km² from the period prior to the major decline in water quality in the 1950s. General linear regression models showed the ecoregion and lake altitude being the best predictors of fish species richness. The number of species was significantly higher in the Alpine than in the Mediterranean ecoregion. Among Alpine lakes, the number of fish species increased significantly with lake volume whilst decreased with altitude. In the Mediterranean lakes, none of the selected parameters was significant. Cluster analysis of fish assemblages (presence/absence) divided the lakes of the Alpine and Mediterranean ecoregions into four and two types, respectively. Pike (Esox lucius), rudd (Scardinius erythrophthalmus) and tench (Tinca tinca) were the main indicator species for the small and mostly shallow lakes in both the Alpine (Type 1) and Mediterranean (Type 6) ecoregions, minnow (Phoxinus phoxinus) for the alpine high altitude lakes (Type 2) and landlocked shad (Alosa fallax lacustris), European whitefish (Coregonus lavaretus) and burbot (Lota lota) for the large and very deep alpine lakes (Type 4). The European whitefish was the only indicator species for the deep Mediterranean lakes (Type 5). These species and associated fish assemblages may be useful indicators in future assessments of the ecological status of Italian lakes according to the European Directives (2000/60/EC and 2008/105/EC).