Some like it deep: Intraspecific niche segregation in ruffe (Gymnocephalus cernua)

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Biogenic methane in freshwater food webs

1. It has long been known that substantial amounts of methane are produced in anoxic lake sediments, and the components of the methane cycle in lakes have been well described. At oxic–anoxic interfaces, methane‐oxidising bacteria (MOB) convert methane to microbial biomass and can be highly productive. However, only recently has methane been recognised as a potentially important carbon and energy source for lake food webs, and some instances have also been reported of methane contribution to river food webs. Stable isotope analysis (SIA) has provided compelling evidence in this respect and has been supplemented by other lines of evidence. 2. In the benthic food webs of lakes, profundal chironomid larvae appear to be the main conduits for trophic transfer of biogenic methane via grazing on MOB. The mode of feeding of these larvae and the microhabitats they generate both promote larval ability to exploit MOB production. Support to chironomid larvae from methane is rather widespread, but its degree is highly variable; estimates suggest that in some lakes methane‐carbon might contribute more than 60% of chironomid carbon biomass. 3. Evidence of crustacean zooplankton in lakes deriving part of their carbon from methane is currently more limited. Reports from some lakes have indicated Daphnia with a substantial (>50%) contribution of methane‐carbon in their biomass. However, for this to happen, an oxic–anoxic interface where sufficient MOB production can occur needs to be within the range of vertical migrations by zooplankton, which may only rarely be the case. Hence, a significant methane subsidy of pelagic food webs in lakes is probably much less widespread than for benthic food webs. 4. There is also recent and currently very limited evidence that some stream benthos derives biomass carbon (reported values up to 30%) from methane. This can occur in stagnant backwater pools where conditions can be analogous to those in lake sediments. However, groundwater aquifers can also supply water supersaturated with methane to some rivers, providing a basis for a microbially‐mediated transfer of methane‐carbon to river benthos. 5. Evidence for significant transfer of methane‐derived carbon to higher trophic levels is still very limited. Within some lakes, those fish species that feed extensively on chironomid larvae can derive a substantial part (perhaps up to 20%) of their carbon biomass from methane. It is also likely that methane‐carbon produced in lakes or rivers is exported to riparian ecosystems when emerging chironomids or other insects are eaten by invertebrate or avian predators. 6. We argue that conceptual models of freshwater food webs, and especially those for lakes, need to be modified to enable incorporation of biogenic methane as a carbon and energy source. For some types of lakes, carbon and energy budgets certainly need to take account of the production and utilisation of biogenic methane, and the accumulating evidence indicates that this is a more widespread phenomenon that has generally been acknowledged hitherto.     

Nutrient release and resuspension generated by ruffe (Gymnocephalus cernuus) and chironomids

1. Benthivorous fish may play an important role in internal nutrient loading. Ruffe are highly specialised, feeding exclusively on bottom animals; thus all nutrients released via their feeding are derived from the bottom and are new to the water column. The fish can also release nutrients from the sediment through resuspension while searching for food. 2. The aim of this study was to estimate experimentally in the laboratory the effect on water quality of resuspension and nutrient release by ruffe and bottom animals (chironomids). 3. Ruffe released nutrients during 8 h experiments as follows: total P 1.4, dissolved PO₄ 0.6, total N 24.0 and NH₄‐N 15.9 μg g^?1 WW h^?1. A decreasing trend in mass‐specific release was observed over time, probably because of starvation. The mass‐specific release of total N and NH₄‐N decreased as the mean weight of fish increased. The mean ratio of excreted N : P was 32. 4. In 26 h experiments with sediment and both ruffe and chironomids, ruffe increased nutrient concentrations and turbidity values significantly but chironomids had an effect only on turbidity. Neither ruffe nor chironomids affected the ratio of inorganic N : P concentrations. An interaction between ruffe and chironomids was found for turbidity. 5. According to these results, benthivorous fish may increase nutrient concentrations in the water column and need to be taken into account when estimating internal loading.     

Microhabitat influence on chironomid community structure and stable isotope signatures in West Greenland lakes

Most functional feeding types are represented within the species rich group of aquatic chironomids. Thus, we hypothesized that different lake types and microhabitats within lakes would (1) host specific chironomid communities and (2) that the individual communities would show specific δ ¹³C stable isotope signatures reflecting the prevailing origin of food source. To test our hypotheses, five lakes in southwest Greenland were investigated at a high taxonomic resolution and with detailed information on δ ¹³C signature of the chironomids and of individual microhabitats (macrophytes, sediment, stones, and profundal). We found that there was a significant difference in δ ¹³C between the chironomid assemblages of freshwater lakes and oligosaline lakes, while assemblages of the littoral microhabitats did not differ significantly. The δ ¹³C of chironomids reflected the wide variety of habitat signals, particularly in the freshwater lakes. Our results indicate that many chironomid taxa are ubiquitous and are found in several microhabitats, suggesting that they can adjust their feeding strategy according to the habitat. The implication is that chironomid assemblage composition has only limited use as indicator of littoral microhabitats in the Arctic. On the other hand, the δ ¹³C signature of fossil chironomids might have a potential as indicator of microhabitats in freshwater lakes.     

Seasonal changes in the stable isotope values of lake-dwelling chironomid larvae in relation to feeding and life cycle variability

1. We studied seasonal changes in the carbon and nitrogen stable isotope ratios of larval Chironomus anthracinus and C. plumosus from the profundal sediments of four contrasting lakes. 2. Pronounced seasonal changes in both δ¹³C and δ¹⁵N values were evident in chironomid larvae of both species from two summer‐stratified, eutrophic lakes: Esthwaite Water and Wyresdale Park. Changes were most marked in the larvae of C. plumosus and in larvae from greater depths. In contrast, neither C. anthracinus in summer‐stratified but mesotrophic Schöhsee, nor C. plumosus in polymictic Großer Binnensee, showed marked seasonality in larval stable isotope ratios. 3. The particularly strong ¹³C‐depletion of larvae from the stratified, eutrophic lakes is attributed to a significant contribution of methane‐derived carbon to their diets. Feeding by larvae on isotopically light methanotrophic bacteria appears to occur mainly when autumn overturn of the water column restores oxygenated conditions to the sediment surface. At this time both δ¹³C and δ¹⁵N values of larvae decreased sharply. 4. Changes in the mean stable isotope ratio of the larval populations can also occur when larger, more isotopically light, larvae pupate and emigrate from the population to hatch as imagos. This effect can induce seasonal changes in larval isotope values even in lakes in which there is no evidence of a significant involvement of methane‐derived carbon in their diets. Variations in emergence patterns between species and between lakes may generate differences in the seasonal pattern of change in stable isotope ratios in larval populations. 5. Our results emphasise the importance of adequate seasonal sampling if stable isotope ratios are to be used as biomarkers to study the role of key groups, such as chironomid larvae, in the trophic structure of lakes.     

Carbon flow in the littoral food web of an oligotrophic lake

Benthic food web dynamics and carbon flow were examined in the littoral zone of Lake Coleridge, a large deep oligotrophic lake, using radioactive and stable isotope techniques in conjunction with analyses of stomach contents of the fauna. We specifically address two hypotheses: (1) that macrophytes only contribute to the carbon flow to higher trophic levels when they have decayed; and (2) that epiphytic algae is the major source of carbon for macroinvertebrates, and thus fish, with only minor contributions from phytoplankton or terrestrial sources. Epiphytic diatoms were a major component of the stomach contents of the gastropod snail Potamopyrgus antipodarum, and of chironomids. Animal remains were also common in the diet of some chironomids, while amorphous organic matter predominated in the stomachs of oligochaetes. A variety of epiphytic algal taxa was found in trichopteran larvae. Feeding rate of P. antipodarum measured with radioactive tracers increased by 10× on decayed macrophytes (Elodea) compared with live material, while feeding rates on characean algae increased by a factor of 3 when decayed material was presented. However, assimilation rates were less than 20% on decayed material compared with 48–52% on live material. Potential carbon sources were easily distinguished based on their ¹³C values, although isotopic ratios showed significant variation among sites. Epiphytic algae showed less variation among sites than macrophytes and were depleted by 4–5 compared with macrophytes. Detrital material, organic matter in the sediments and plankton were significantly depleted in ¹³C relative to macrophytes and slightly depleted relative to epiphytic algae. Most macroinvertebrate taxa showed a similar pattern among sites to macrophytes and epiphytic algae. P. antipodarum and chironomids were slightly enriched compared with epiphytic algae. Ratios for the common bully (Gobiomorphus cotidianus) were generally consistent with a diet dominated by chironomids, while there was some evidence for terrestrial inputs for koaro (Galaxias brevipinnis) and juvenile brown trout. Epiphytic algae appear to underpin much of the production in the littoral zone of this oligotrophic lake, with trichopteran and chironomid larvae mediating carbon flows from algae to fish. Macrophytes do not make a major contribution directly to carbon flow to higher trophic levels even when decayed. The lack of a direct link between macrophytes and higher trophic levels is due to the faunal composition, including a lack of large herbivores.     

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.

     

Food and habitat of four Xenotilapia species (Teleostei, Cichlidae) in a sandy bay of northern Lake Tanganyika (Burundi)

The feeding ecology of four species of Xenotilapia (Teleostei, Cichlidae) from Lake Tanganyika was investigated by examining their intestine contents in three different habitats from two 24-h catch cycles. The fish were caught in a sandy bay 4 km north-west of Bujumbura (Burundi) during the dry season, from August until October 1993. Three different habitats were sampled: an area 600 m offshore at a depth of 15 m, a littoral zone near a reed bed (depth, 0–5 m), and an area near a sandy beach (depth, 5 m). The fish were most active in the early morning and at night, except for X. caudafasciata which was the most diurnal. X. ochrogenys was confined to the vicinity of the reeds, where few individuals of the other species were found, and prey comprised mainly ostracods, copepods and chironomid larvae (Tanytarsini). X. ornatipinnis and X. longispinis occurred in large numbers both at a depth of 15 m and near the sandy beach. X. ornatipinnis preyed mainly on copepods, oligochaetes, diatoms and chironomid larvae. X. longispinis had a wider range of temporal and spatial distribution compared with the other species and fed on copepods, ostracods and chironomid larvae (mainly Microchironomus spp.). X. caudafasciata (only caught at 15 m) fed mostly on profundal chironomid larvae (Tanypodinae and Microchironomus spp.). Trophic niche overlap between X. longispinis and X. caudafasciata appeared to be reduced through different activity patterns. Few intraspecific differences in food diet were found between sex and age groups.     

Depth gradients in food‐web processes linking habitats in large lakes: Lake Superior as an exemplar ecosystem

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Carbon sources for lake food webs in the Canadian High Arctic and other regions of Arctic North America

We investigated the role of autochthonous and terrestrial carbon in supporting aquatic food webs in the Canadian High Arctic by determining the diet of the dominant primary consumer, aquatic chironomids. These organisms were studied in fresh waters on 3 islands of the Arctic Archipelago (~74–76°N) including barren polar desert watersheds and a polar oasis with lush meadows. Stomach content analysis of 578 larvae indicated that chironomids primarily ingested diatoms and sediment detritus with little variation among most genera. Carbon and nitrogen stable isotope mixing models applied to 2 lakes indicated that benthic algae contributed 68–95% to chironomid diet at a polar desert site and 70–78% at a polar oasis site. Detritus, originating from either phytoplankton or terrestrial sources, also contributed minor amounts to chironomid diet (0–32%). Radiocarbon measurements for the 2 lakes showed that old terrestrial carbon did not support chironomid production. Carbon stable isotope ratios of chironomids in other High Arctic lakes provided further dietary evidence that was consistent with mixing model results. These findings indicate that, in the Canadian High Arctic, chironomids (and fish that consume them) are supported primarily by benthic algae in both polar desert and oasis lakes. In contrast, our review of carbon flow studies for lakes in other Arctic regions of North America shows that terrestrial carbon and phytoplankton can be important energy sources for consumers. This study provides a baseline to detect future climate-related impacts on carbon pathways in High Arctic lakes.     

Diel-feeding activity in early summer of racer goby Neogobius gymnotrachelus (Gobiidae): a new invader in the Baltic basin

The first recording of the Ponto‐Caspian racer goby in Poland was during 1995 in the River Bug (River Vistula system). Within 5 years, the species had spread to the downstream section of the Vistula. One of the potential impacts of invasive species on native fauna is competition for food. Therefore, the diel patterns in diet composition and gut fullness coefficient (FC) of racer goby were examined at one study site in the W?oc?awski Reservoir (lower River Vistula), during May 2003. An average of 20 individuals were examined each 4 h over one 24‐h period (125 fish in total). The proportion of main food items and diet width did not differ among three size‐groups (43–59, 60–79 and 80–97 mm total length), and the relative biomass ratio of main food categories did not differ over the diel cycle. Amphipods constituted 11–70% of total gut content biomass and were found on average in 84% of analysed alimentary tracts. The second prey types were chironomid larvae (16–63% of total food biomass; frequency occurrence: 61–91%), and to a lesser extent chironomid pupa, ceratopogonid larvae, oligochaets, dipteran imagines and copepods, with fish larvae found in the gut of eight gobies. Gut fullness coefficient (FC) differed significantly over the 24‐h period, with the highest value at night and in early morning. In conclusion, racer goby forages mainly on benthos and has a nocturnal‐feeding activity. No significant diet overlap was found between racer goby and native percids, i.e. Eurasian perch Perca fluviatilis and ruffe Gymnocephalus cernuus.     

Biogenic methane contributes to the food web of a large,shallow lake

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Chironomid communities as water quality indicators

Recent mathematical indices summarizing biological communities of indicators are recapitulated. Improvements of these indices based on weighting according to width of trophic ranges of each species are suggested. Their principle deficiencies, however, are pointed out.
Revised lists of characteristic profundal as well as littoral and sublittoral chironomids in Nearctic and Palearctic lakes show that at least 15 characteristic chironomid species communities can be delineated, 6 in each of the oligotrophic and the eutrophic ranges and 3 in the mesotrophic range. It is proposed that these communities be lettered consecutively in the Greek alphabet from α (alpha) to o (omikron). A key to the 15 divisions based on the species associations in the profundal zone of harmonic lakes is put forward. There is very good correlation between the 15 divisions and the ratios of average total phosphorus to mean lake depth and average chlorophyll a to mean lake depth.
The ratio of chironomids to oligochaetes and the distribution patterns of single species have proven useful in pin-pointing localized areas of pollution. The primary mechanism governing the distribution of chironomid communities in oligotrophic and mesotrophic lakes appears to be the availability of food materials rather than the annual hypolimnetic oxygen concentration. In eutrophic lakes the relationships between organic matter accumulation and oxygen levels are so interdependent as to be inseparable.     

Influences of slimy sculpin (Cottus cognatus) predation on the rocky littoral invertebrate community in an arctic lake

We tested the role of the slimy sculpin (Cottus cognatus), a benthic fish, in structuring the rocky littoral invertebrate community in Toolik Lake, Alaska. Comparisons of sculpin gut contents and prey community structure indicated that these fish forage selectively, eating proportionally more large and motile prey, and proportionally fewer small and sessile forms. Field experiments compared the effects of natural, reduced and elevated sculpin densities on benthic community structure. At natural levels of sculpin density, biomass of trichopteran larvae were reduced by more than 50%, and predatory chironomid larvae by 27%, in comparison to areas where sculpin were excluded. Tube-dwelling and small free living chironomid larvae were unaffected at normal sculpin densities. Under artificially high sculpin densities, there was some reduction of tube-dwelling chironomids, but the small free living ones remained unaffected. There appears to be a threshold length of about 3.5 mm, below which chironomid larvae are free form sculpin predation. Tube-dwelling chironomids may be longer than this threshold, but still avoid predation by having most of their body hidden in their tubes.     

Rapid changes in fish community structure and habitat distribution following the precipitation of lake phosphorus with aluminium

1. Fish community structure and habitat distribution of the abundant species roach, perch and ruffe were studied in Lake Nordborg (Denmark) before (August 2006) and after (August 2007) aluminium treatment to reduce internal phosphorus loading. 2. Rapid changes in fish community structure, abundance and habitat distribution occurred following a decline in in‐lake phosphorus concentrations from 280 to 37 μg P L^?1 and an increase in Secchi depth transparency from 1.1 to 1.9 m (August). The proportion of perch in overnight gill net catches increased, whilst roach decreased, and the average weight of all key species increased. 3. The habitat distribution of perch and roach changed from a high proportion in the upper pelagic and littoral zones in 2006, towards enhanced proportions in the deeper pelagic and profundal zone in 2007. The abundance of large‐bodied zooplankton increased and the abundance of benthic invertebrates decreased in the same period, suggesting that the habitat shift was not induced by food limitation. 4. Ruffe shifted from the littoral and upper profundal zones towards the deep profundal zone, likely reflecting an increased predation risk in the littoral zone and better oxygen conditions in the deep profundal. 5. Our results indicate that enhanced risk of predation in the upper pelagic and the littoral zones and perhaps improved oxygen concentrations in the deeper profundal zone at decreasing turbidity are responsible for the observed habitat shift. The results indicate that fish respond rapidly to changes in nutrient state, both in terms of community structure and habitat use.     

Temporal and spatial distribution of chironomid larvae and oligochaetes in two Dutch storage reservoirs

Long-term changes in distribution and taxonomic composition of chironomid larvae and oligochaetes in two water storage reservoirs in The Netherlands were studied. A succession among the chironomid species was observed. In the last 10–15 years chironomid densities varied. Compared with other lakes chironomid densities were high. Densities of chironomids were higher in the littoral zone than in the profundal zone. The opposite was found for oligochaetes. Densities and composition of the chironomid fauna in the two reservoirs were similar.Procladius, Tanytarsus andHarnischia dominated at all depths. However, pupal exuviae samples showed some differences in chironomid taxonomic composition between the two reservoirs. Orthocladiinae, rarely found in bottom samples, abounded in pupal exuviae samples.     

Reconstructing long‐term changes (150 years) in the carbon cycle of a clear‐water lake based on the stable carbon isotope composition (δ13C) of chironomid and cladoceran subfossil remains

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Incipient speciation through niche expansion: an example from the Arctic charr in a subarctic lake

Two reproductive isolated morphs of Arctic charr (Salvelinus alpinus), termed profundal and littoral charr according to their different spawning habitats, co-occur in the postglacial lake Fjellfr?svatn in North Norway. All profundal charr live in deep water their entire life and have a maximum size of 14cm, while the littoral charr grow to 40cm. Some small and young littoral charr move to the profundal zone in an ontogenetic habitat shift in the ice-free season and the rest of the population remains in epilimnic waters. The two morphs had different diet niches in the profundal zone: the profundal charr ate typical soft-bottom prey (chironomid larvae, pea mussels and benthic copepods), while the young littoral charr mainly consumed crustacean zooplankton. In four other lakes without a profundal morph (i.e. monomorphic populations), young charr also performed ontogenetic habitat shifts to the profundal zone and fed on zooplankton. The profundal morph of Fjellfr?svatn therefore utilize a food resource niche that neither the littoral morph nor comparable monomorphic populations exploit. This suggests that intraspecific resource competition has driven incipient ecological speciation of the profundal charr of Fjellfr?svatn. The exploitation of the soft-bottom resources by the profundal charr supports earlier experimental findings that the profundal morph is genetically different in trophic behaviour and morphology. The sympatric ecological divergence within the profundal habitat is possible because unexploited food resources (soft-bottom profundal prey) are available. Apparently, this represents a case of incipient segregation by expansion to new resource types (niche invasion), and not by subdivision of one broad ancestral niche.     

Habitat, diet and food assimilation of Arctic charr under the winter ice in two subarctic lakes

The Arctic charr Salvelinus alpinus populations of the subarctic lakes Takvatn and Fjellfrøsvatn, north Norway, concentrated in the littoral zones (0–15 m) of the lakes during the entire winter (December to May) despite very low temperatures (0·2 and 0·7° C). High prey availability, low predation and competition and comparatively better light under snow and ice in shallow compared with deep water are probable reasons. At ice break in June, all Arctic charr moved to the profundal zone for a brief period, probably in response to the sudden light increase and a profundal resource peak of chironomid pupae. In the summer, the Arctic charr are found in the pelagic, profundal and littoral zones of the lakes. These populations therefore perform regular habitat shifts between the littoral zone in the winter, the profundal zone at ice break and the whole lake in the summer and autumn. The fish fed continuously during winter despite the cold water and the poor light. Amphipods and chironomid larvae dominated the diet. Catch per unit effort, numbers of stomachs with food and food intake rates varied with the subarctic light cycle but were lowest after the winter solstice. The winter assimilation of energy was about equal to the standard metabolism in Takvatn but was higher in Fjellfrøsvatn. The assimilation increased in both lakes under the spring ice in May. The habitat choice, diet and energy assimilation indicate that the Arctic charr is well adapted to the extreme winter conditions of subarctic lakes.     

Trophic level assessment of profundal sediments of the artificial Lake Campotosto (Central Italy), using midge larval community (Diptera: Chironomidae)

A study on the profundal chironomids of the artificial Lake Campotosto (Central Italy) was carried out during the summer/early autumn of 1983 and 1984, in order to analyse their composition and community structure in relation to the lake trophic level assessed by water chemical analysis.A total of about 2000 specimens belonging to 15 taxa were collected during the study.Chironomus plumosus group andTanytarsus spp. dominated in 1983 and 1984, respectively, showing a competitive relationship probably due to the larval size. The functional feeding organization was mostly composed of collectors (percentages greater than 90%), revealing the presence of abundant fine organic deposits (FPOM). Diversity and evenness appeared to be negatively affected by the monotony of food, which seems to constitute the key factor in governing both the taxonomic and the trophic structure of chironomid fauna.A clear discrepancy between water chemical data and profundal chironomid analyses was observed in the assessment of the lake trophic level. Sediments exhibited eutrophicated conditions, whereas overlying waters indicated an oligotrophic status. The relevance of profundal macrobenthic investigations in detecting eutrophication is stressed.