A trophic pathway from biogenic methane supports fish biomass in a temperate lake ecosystem

Although some primary consumers such as chironomid larvae are known to exploit methane‐derived carbon via microbial consortia within aquatic food webs, few studies have traced the onward transfer of such carbon to their predators. The ruffe Gymnocephalus cernuus is a widespread benthivorous fish which feeds predominantly on chironomid larvae and is well adapted for foraging at lower depths than other percids. Therefore, any transfer of methanogenic carbon to higher trophic levels might be particularly evident in ruffe. We sampled ruffe and chironomid larvae from the littoral, sub‐littoral and profundal areas of Jyväsjärvi, Finland, a lake which has previously been shown to contain chironomid larvae exhibiting the very low stable carbon isotope ratios indicative of methane exploitation. A combination of fish gut content examination and stable isotope analysis was used to determine trophic linkages between fish and their putative prey. Irrespective of the depth from which the ruffe were caught, their diet was dominated by chironomids and pupae although the proportions of taxa changed. Zooplankton made a negligible contribution to ruffe diet. A progressive decrease in δ¹³C and δ¹⁵N values with increasing water column depth was observed for both chironomid larvae and ruffe, but not for other species of benthivorous fish. Furthermore, ruffe feeding at greater depths were significantly larger than those feeding in the littoral, suggesting an ontogenetic shift in habitat use, rather than diet, as chironomids remained the predominant prey item. The outputs from isotope mixing models suggested that the incorporation of methane‐derived carbon to larval chironomid biomass through feeding on methanotrophic bacteria increased at greater depth, varying from 0% in the littoral to 28% in the profundal. Using these outputs and the proportions of littoral, sub‐littoral or profundal chironomids contributing to ruffe biomass, we estimated that 17% of ruffe biomass in this lake was ultimately derived from chemoautotrophic sources. Methanogenic carbon thus supports considerable production of higher trophic levels in lakes.     

Factors governing the spatial and temporal distribution of Chironomid larvae in the Maarsseveen lakes with special emphasis on the role of oxygen conditions

Distribution patterns of the larvae of Chironomidae are compared in three water systems in The Netherlands, which vary in trophic state and oxygen regimes. The life cycles and flying periods of some dominant chironomid species in two of the investigated lakes, Lakes Maarsseveen I and II, are determined by comparing data on the seasonal variations in larval densities with existing literature on Chironomidae in the Maarsseveen lakes. In the oligo-mesotrophic Lake Maarsseveen I (LM I), hypoxic or anoxic conditions in the hypolimnion are observed only at the end of the stratification period. A clear zonation of the chironomid fauna is present in this lake. The littoral zone is dominated byCladotanytarsus gr.mancus andStictochironomus sticticus, the littoriprofundal zone byTanytarsus bathophilus, and the profundal zone byChironomus anthracinus. In comparison with the other species in LM I,T. bathophilus larvae show the most variable distribution patterns over time. Larvae are found in all depths from July to September, but disappear from the hypolimnion as soon as oxygen conditions deteriorate. In the eutrophic Lake Maarsseveen II (LM II), oxygen depletion of the hypolimnion starts immediately after the onset of the thermal stratification in June, and continues until autumnal turnover in November. In this lake, the chironomid community consists primarily ofS. sticticus andCl. gr.mancus, and is confined to the narrow littoral zone. No chironomid larvae are found in the deeper parts of the lake. The eutrophic Lake Gijster in the Brabantse Biesbosch is a deep, man-made reservoir, that is artificially destratified during the summer. In this lakeTanytarsus bathophilus is found in the profundal sediments, whereas almost noChironomus is found in this zone. It is concluded that oxygen conditions existing in the deeper regions of the investigated lakes in large part determine the occurrence and distribution of chironomid species. The distribution ofT. bathophilus is limited by unfavorable oxygen conditions and not by the trophic state of the lake. These findings are part of a thesis (HEINIS, 1993).     

Phenology and bathymetric distribution of the profundal chironomid fauna in Starnberger See (F.R. Germany)

Results of the first long-term investigation of the profundal chironomid fauna of Starnberger See, a prealpine lake are reported. Data are presented on the qualitative and quantitative composition of the chironomid fauna in the profundal of Starnberger See, on the spatial variation, and on the phenology of the dominant species, based on pupal occurrence and the seasonal distribution of larval instars. Twenty-five distinct chironomid taxa could be recorded in the profundal zone. Typical inhabitants of the profundal zone of Starnberger See are the larvae of the genus Micropsectra. The maximum abundance of Micropsectra larvae were found at 60 m with 5644 Ind.m⁻². Abundance, distribution and phenology of the dominant chironomid species are discussed in comparison with other lakes.     

Distribution of chironomids (Diptera) in low arctic West Greenland lakes: trophic conditions, temperature and environmental reconstruction

1. Surface sediment samples of subfossil chironomid head capsules from 47 lakes in southern West Greenland were analysed using multivariate numerical methods in order to explore the relationship between chironomid assemblages and selected environmental variables. The study lakes are located along a climate gradient ranging from coastal maritime conditions near the Davis Strait to a continental climate near the margin of the Greenland ice sheet. 2. High‐resolution surface water temperatures were measured through the summer season using automatic data loggers in 21 of the study lakes. The mean July surface water temperature (1999) ranged from 7.3 to 16.5 °C in the data set. 3. In all lakes, a total of 24 chironomid taxa were recorded; Micropsectra, Psectrocladius, Chironomus and Procladius were the dominant genera. There was a strong correlation between the trophic variables [total nitrogen and total phosphorus (TN, TP)] and temperature, and in redundancy analysis (RDA) the three variables explained almost equal significant amounts of variation in the chironomid data (19.8–22.3%). However, temperature lost significant explanatory power when the effect of TN was partialled out in RDA. 4. The lakes were classified using two‐way indicator species analysis (TWINSPAN ) into eight groups defined by temperature, trophic variables, salinity (conductivity) and lake‐morphometric data. Fourteen chironomid taxa showed significant differences in percentage abundances among groups, with Heterotrissocladius, Micropsectra, Ablabesmyia and Chironomus as the most robust group‐indicator taxa. Forward selection of taxa in multiple discriminant analysis was used to fit chironomid assemblages into lake groups. Using only eight taxa, 95% of lakes were correctly classified at a second TWINSPAN division level (four groups) and 85% of lakes at a third division level (eight groups). 5. This study showed that there is considerable potential in using subfossil chironomid head capsules as paleoenvironmental indicators in both short‐ and long‐term (down‐core) studies of lake ontogeny and palaeoclimate conditions in West Greenland. However, because of the strong correlation between temperature and trophic variables, a quantitative reconstruction of lake‐ and habitat‐type is recommended, in combination with direct reconstruction of single variables such as temperature.     

Subfossil chironomid assemblages in deep,stratified European lakes: relationships with temperature,trophic state and oxygen

1. The distributions of subfossil remains of chironomid larvae in 28 large, deep and stratified lakes in Europe were examined in surface sediments along a latitudinal transect ranging from northern Sweden to southern Italy. 2. Canonical correspondence analysis (CCA) showed that summer surface water and July air temperature, as well as total phosphorus (TP) concentrations, hypolimnetic oxygen availability and conductivity were statistically significant (P < 0.05) explanatory variables explaining between 11 and 14% of the variance in the chironomid data. 3. Owing to the spatial scale covered by our study, many environmental variables were covarying. Temperature, TP concentration and oxygen availability were positively or negatively correlated with the first axis of a detrended correspondence analysis (DCA) of chironomid assemblages, suggesting that climatic and trophic conditions influenced profundal chironomid assemblages either in a direct (food and oxygen) or in an indirect (temperature) way. Parameters related to local environmental conditions, lake morphology and bedrock geology, such as organic matter content of the sediment, maximum lake depth, Secchi depth and pH, were not significant in explaining the distribution of chironomid assemblages in our study lakes. 4. The strong relationship between chironomid assemblages and summer temperature may be related to the covariation of temperature with parameters, such as nutrient and oxygen availability, known to affect chironomid assemblages in deep, stratified lakes. However, summer temperature explained a statistically significant proportion of the variance in the chironomid assemblages even when effects of oxygen availability and TP concentrations were partialled out. This suggests that summer temperature has an effect on chironomid assemblages in deep lakes, which is not related to its covariation with trophic state. 5. The potential of fossil chironomid analysis for quantitatively reconstructing past nutrient conditions in deep, stratified lakes was examined by calculating the Benthic Quality Index (BQI) based on subfossil chironomids and by comparing BQI values with observed TP concentrations. BQI was linearly related to log‐transformed TP. Applying this relationship to fossil chironomid assemblages from Lake Päijänne (Finland) produced a TP reconstruction in agreement with measured TP during the period 1970–1990, demonstrating that this approach can provide quantitative estimates of past nutrient concentrations in deep, stratified lakes.     

Assessing lake eutrophication using chironomids: understanding the nature of community response in different lake types

1. Total phosphorus (TP) and chlorophyll a (Chl a) chironomid inference models ( Brodersen & Lindegaard, 1999 ; Brooks, Bennion & Birks, 2001 ) were used in an attempt to reconstruct changes in nutrients from three very different lake types. Both training sets were expanded, particularly at the low end of the nutrient gradient, using contemporary chironomid assemblages and environmental parameters from 12 British lakes, although this had little improvement on the model performances. 2. Dissimilarity analyses showed that the historic chironomid assemblages did not have good analogues in the original calibration or extended datasets. However, since the transfer functions are based on weighted averages of the trophic optima for the taxa present and not on community similarities, reasonable downcore inferences were produced. Ordination analyses also showed that the lakes retain their ‘identity’ over time, as the sample dissimilarities within lakes were less than the dissimilarities between lakes. 3. Analysis of the three historic lake profiles showed a range of chironomid community responses to lake development. Chironomids from a shallow lake, Slapton Ley, responded indirectly to nutrient enrichment (TP), probably through altered substrate, macrophyte and fish conditions, rather than directly to primary productivity (Chl a). A stratified lake, Old Mill Reservoir, showed a loss of the profundal chironomid fauna due to increasing primary productivity (Chl a) coupled with increasing hypoxia. A response to nutrients (TP or total nitrogen (TN)) at this site is also indirect, and the TP reconstruction therefore cannot be reliably interpreted. The third lake, March Ghyll Reservoir has little change in historic chironomid communities, suggesting that this well mixed, relatively unproductive lake has changed less than the other lakes. 4. Using chironomids to reconstruct nutrient histories does not follow a simple scheme. The response to changes in nutrients may be direct, but mediated through other ecosystem components. As alternative stable states are possible at a given level of TP it is also likely that alternative chironomid communities exist under similar nutrient conditions. Changes in biological communities can thus occur over thresholds, and it is only biological proxies that can reflect such ecosystem switches within palaeoenvironmental investigations.     

Can lake restoration by fish removal improve the status of profundal macroinvertebrate assemblages?

相似文献   

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

相似文献   

Responses of insects,especially Chironomidae (Diptera), and mites to 130 years of acidification in a Scottish lake

About 130 years of anthropogenic acidification of Round Loch of Glenhead, SW Scotland, has resulted in successively decreased stability, diversity, productivity and survival rate of the non-biting midge (Diptera: Chironomidae) fauna. Similar trends have also been observed among mayflies, caddis-flies and water mites. The first effects of anthropogenic acidification on the insect and mite fauna, as evidenced by palaeolimnological analyses of²¹⁰Pb-dated sediment cores, occurred as early as around 1850, i.e. earlier than in any other lake hitherto studied. The drop in the lake's pH was first indicated by decreased stability and changes in species composition of chironomids and mayflies in the littoral zone. Major changes in the profundal chironomid fauna did not take place until about 1950, when mean lake pH dropped below 5.0. At the same time, the littoral insect fauna became even more unstable and the first significant elimination of species occurred. Comparison with insect fauna of other lakes suggests that a pH of less than 5 might be critical for the ecological conditions in many acidified lakes. None of the twelve most common chironomid species present prior to the acidification of the lake had disappeared after 120–130 years of considerable acidification, and they are all common in oligotrophic lakes with a pH of 6.5–7.0. This is in contrast to the effects of heavy acidification on other aquatic animal groups. Chironomids are probably more sensitive to lake trophic status than lake acidity.     

Disappearance of deep profundal zoobenthos in Lake Ikeda, southern Kyushu, Japan, with relation to recent environmental changes in the lake

A benthological survey in a deep caldera, Lake Ikeda, southern Kyushu, Japan, in 1998 revealed that no zoobenthos were found in the deep profundal, although two tubificid oligochaetes, Tubifex tubifex and Limnodrilus hoffmeisteri, and a chironomid, Procladius sp., were distributed in the upper profundal zone. This is the first record of oligochaete composition in the lake. Lake Ikeda had been typically oligotrophic until the 1940s, and zoobenthic assemblages were recorded throughout the profundal bottom in the 1920s and 1970s. Recent disappearance of the deep profundal zoobenthos could be caused by the stagnation of anoxic waters in the hypolimnion, in connection with eutrophication triggered by nutrient loading, as well as change in the thermal circulation system presumably caused by global warming.     

Changes in fossil chironomid remains along a depth gradient: evidence for common faunal thresholds within lakes

Many environmental variables that are important for the development of chironomid larvae (such as water temperature, oxygen availability, and food quantity) are related to water depth, and a statistically strong relationship between chironomid distribution and water depth is therefore expected. This study focuses on the distribution of fossil chironomids in seven shallow lakes and one deep lake from the Plymouth Aquifer (Massachusetts, USA) and aims to assess the influence of water depth on chironomid assemblages within a lake. Multiple samples were taken per lake in order to study the distribution of fossil chironomid head capsules within a lake. Within each lake, the chironomid assemblages are diverse and the changes that are seen in the assemblages are strongly related to changes in water depth. Several thresholds (i.e., where species turnover abruptly changes) are identified in the assemblages, and most lakes show abrupt changes at about 1–2 and 5–7 m water depth. In the deep lake, changes also occur at 9.6 and 15 m depth. The distribution of many individual taxa is significantly correlated to water depth, and we show that the identification of different taxa within the genus Tanytarsus is important because different morphotypes show different responses to water depth. We conclude that the chironomid fauna is sensitive to changes in lake level, indicating that fossil chironomid assemblages can be used as a tool for quantitative reconstruction of lake level changes.     

Allometric trajectories of body and head morphology in three sympatric Arctic charr (Salvelinus alpinus (L.)) morphs

A study of body and head development in three sympatric reproductively isolated Arctic charr (Salvelinus alpinus (L.)) morphs from a subarctic lake (Skogsfjordvatn, northern Norway) revealed allometric trajectories that resulted in morphological differences. The three morphs were ecologically assigned to a littoral omnivore, a profundal benthivore and a profundal piscivore, and this was confirmed by genetic analyses (microsatellites). Principal component analysis was used to identify the variables responsible for most of the morphological variation of the body and head shape. The littoral omnivore and the profundal piscivore morph had convergent allometric trajectories for the most important head shape variables, developing bigger mouths and relatively smaller eyes with increasing head size. The two profundal morphs shared common trajectories for the variables explaining most of the body and head shape variation, namely head size relative to body size, placement of the dorsal and pelvic fins, eye size and mouth size. In contrast, the littoral omnivore and the profundal benthivore morphs were not on common allometric trajectories for any of the examined variables. The findings suggest that different selective pressures could have been working on traits related to their trophic niche such as habitat and diet utilization of the three morphs, with the two profundal morphs experiencing almost identical environmental conditions.     

The impact of acidification on Chironomidae (Diptera) as indicated by subfossil stratification

The impact of acidification on the chironomid profundal fauna was studied by comparing chironomid head capsule deposits from different sediment layers. The 14–15 cm level was regarded as relatively unaffected by acid precipitation and therefore used as a control. The total number of capsules decreased, especially the number of Tanytarsinii capsules, from deeper to more superficial layers. Phaenopsectra and Psectrocladius showed a relative increase towards the surface level of the sediment. Probable causes for these changes are discussed.     

Tracing lake trophic history with a chironomid–total phosphorus inference model

1. In the absence of historical water chemistry data, predictive biological indicator groups preserved in lake sediments can be employed to reconstruct the history of lake eutrophication. Diatoms are well established in this role, but to augment diatom‐based inferences of nutrient status we investigate the potential use of chironomid midges (Insecta: Chironomidae). 2. Canonical correspondence analysis (CCA) of modern chironomid assemblages in surface sediments from 44 lakes in the English Midlands and Wales, U.K., shows that five environmental variables (total phosphorus (TP), bottom dissolved oxygen, maximum lake depth, Secchi depth and surface water temperature) make a statistically significant (P < 0.05) contribution to explaining the variance in the chironomid data, of which TP makes the largest contribution (29%). 3. The relationship is used to develop a series of weighted averaging (WA) and partial least squares (PLS), (WA‐PLS) models to infer log₁₀TP. The models are evaluated by leave‐one‐out (jack‐knifing) cross‐validation. The simplest minimal adequate model is provided by WA with unweighted inverse deshrinking of root mean square error of prediction (RMSEP_jack=0.34 and r²_jack=0.60). 4. Using this model, the trophic history of Betton Pool, Shropshire, U.K., is reconstructed from the mid‐19th century to the present day and the results from the chironomid‐TP model are compared with inferences from a diatom‐TP model ( 13 ). Both reconstructions suggest that there was a gradual rise in TP since 1850 AD until about 1974, followed by a more pronounced and rapid increase that has continued until the present. Inferred TP values from the WA chironomid inference model agree with diatom‐inferred values. 5. The study demonstrates that fossil chironomid assemblages can be used to investigate quantitatively the trophic history of lakes.     

The role of chironomids (Diptera) in the profundal macrozoobenthos in Finnish reservoirs

The proundal macrozoobenthos of 12 Finnish inland polyhumic reservoirs was studied by quantitative sampling in 1984 and 1986. Additionally, the succession of the macrozoobenthos in one young reservoir (Lake Kyrkösjärvi) was studied during its first six years. The zoobenthos material was analyzed using the DCA (detrended correspondence analysis) method, and the ordination of sites (i.e. reservoirs), was related to associated environmental data. The role of chironomids was, in general, important in the profundal of the reservoirs. Because of the coarse bottom quality at many sampling sites, many littoral taxa were found in the profundal zone. The genusChironomus dominated (>50% of the total zoobenthic biomass) especially in the acidic, upper-reach reservoirs.Chironomus larvae also dominated in the zoobenthos of Lake Kyrkösjärvi during its first three years. Some of the reservoirs showed similarities with the dystrophic lake type of the lake typology system (Brundin, 1956). Because of the initially eutrophic character of the bottom, many reservoirs belonged to eutrophic lake types. Finnish reservoirs appear to contain a zoobenthic fauna intermediate between natural lakes and bog pools. This theme was discussed from the ecological and evolutionary point of view ofChironomus species.     

Predictors of Seasonal Oxygen Levels in Small Florida Lakes: The Importance of Color

This study examines the relationship of profundal oxygen concentrations in 55 shallow Florida lakes to humic color, trophic state, and lake size during different seasons. The data set represented a broad range of color and trophic state. The percent saturation of dissolved oxygen remained relatively constant during the fall (mean 78.4%), winter (mean 81.3%), and spring (mean 82.5%), but declined markedly during summer (mean 65.2%). Chlorophyll a concentrations were highest during the winter (mean 2.52 mg m^–3) and lowest during the fall (mean 1.17 mg m^–3), while color peaked during the fall (mean 30.1 mg Pt l^–1) and was lowest during the summer (mean 12.7 mg Pt l^–1). The relative importance of lake size, chlorophyll a, and color in explaining variation in percent oxygen saturation was examined using multiple regression. Percent oxygen saturation was negatively correlated with color during the winter, spring, and summer, and positively correlated with lake size in the winter and spring. However, percent oxygen saturation showed no relationship with chlorophyll a during any season. These results suggest that colored Florida lakes are naturally oxygen depleted and that profundal oxygen values have little relationship to lake trophic state. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

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.     

Bottom-up effects of bream (Abramis brama L.) in Lake Balaton

Enclosures (17 m³) were used in the mesotrophic area of Lake Balaton to determine the impact of benthivorous bream (Abramis brama L.) on the lower trophic levels during summers of 1984–86. In enclosures with a fish biomass similar to the biomass in the eutrophic area of the lake, the number of phytoplankton species was highest. In enclosures with a low fish biomass the phytoplankton was dominated by the greens. A high biomass of bream in the mesotrophic basin caused bacterial production corresponding to that of the eutrophic part of the lake. Crustaceans were dominated by copepods and were unable to control phytoplankton peaks. Bottom-up effects of bream were more obvious than top-down effects and seem to be more important in the possible control of water quality.     

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.