Influence of Chironomidae (Diptera) faecal pellet accumulation on lake sediment quality and larval abundance of pestiferous midge Glyptotendipes paripes

Two opposite distribution patterns of larval Glyptotendipes paripes in relation to organic carbon content in sediments of central Florida lakes were discovered. In a majority of examined lakes, G. paripes larvae were most abundant in sand sediment and their density rapidly declined with increased carbon content (type 1 lakes); however, in some cases the opposite was true (type 2 lakes). To elucidate this anomaly, field-collected organic sediments from types 1 and 2 lakes and sand sediment were studied for G. paripes development in the laboratory. Type 1 organic sediment consisted predominantly of fine particles (<0.25 mm diameter) with low dissolved oxygen levels, whereas type 2 organic sediments consisted primarily of chironomid large faecal pellet aggregates (>0.25 mm diameter), with significantly higher levels of dissolved oxygen concentrations that were similar to sand sediment. Type 2 organic sediment and sand sediment were conducive to higher survival of G. paripes larvae than fine organic sediment. The larvae in type 2 organic sediment produced longer tubes than in other sediment types. This observation indicates that accumulation of chironomid faecal pellets in lake sediments may change physical properties, such as dissolved oxygen level and consequently alter conditions for survival of chironomid larvae and possibly other benthic fauna.     

A spatially explicit computer model for immature distributions of Glyptotendipes paripes (Diptera: Chironomidae) in central Florida lakes

Density and spatial distribution data of immature Glyptotendipes paripes Edwards (Diptera: Chironomidae), in relation to selected water and sediment characteristics prevailing in three central Florida lakes, were used to develop and calibrate a qualitative model of G. paripes distributions with the purpose of developing more efficient sampling plans for research or population management. For the model, each lake habitat was defined by a two-dimensional matrix, while a three-dimensional matrix was used to simulate the life stage-structured population. A Lefkovitch population matrix was used to project survival and development of the population in each spatial unit. The model incorporated lake bathymetry, sediment dry weight, water level, food availability (as a function of Secchi disk transparency) and strength of density dependence as influences on immature survival and adult fecundity. Temperature-dependent development data for G. paripes were incorporated to estimate development time. A simple redistribution function was used to simulate the dispersal of adults over the simulated habitat for oviposition. Immatures and relevant environmental data taken for eight dates, at each of two other central Florida lakes were used to validate the model. The mean correct prediction rate of the model for field density distributions within 0.5 log(n+1) immatures/m² density in spatial habitat cells was 0.64 and 0.66 for the validation lakes. Presence/absence correct prediction rates were 0.61 and 0.76, while matching to a proposed sampling stratification based on a treatment threshold was 0.85 and 0.88. These values indicate that the model is efficient for preparing stratified immature G. paripes sampling plans for central Florida lakes.     

The impact of Chironomus plumosus larvae on organic matter decay and nutrient (N, P) exchange in a shallow eutrophic lake sediment following a phytoplankton sedimentation

The importance of Chironomus plumosus larvae onbenthic metabolism and nutrient exchange across thesediment–water interface was evaluated in a shalloweutrophic lake (Lake Arreskov, Denmark) following aphytoplankton sedimentation. Chironomus plumosuslarvae were added to laboratory sediment microcosms,corresponding to a density of 2825 larvae m⁻².Non-inhabited microcosms served as controls. Asedimentation pulse of organic matter was simulated byadding fresh algal material (Chlamydomonasreinhardii) to sediment cores (36 g dryweight m⁻²). The mineralization was followed bymeasuring fluxes of O₂, CO₂, dissolvedinorganic nitrogen and phosphate. A rapid clearance ofalgae from the water column in faunated microcosmssuggested that chironomids may be of major importancein controlling phytoplankton concentrations in shalloweutrophic lakes. Chironomids increased the sedimentO₂ uptake ≈ 3 times more than what wouldbe expected from their own respiration, indicating astimulation of microbial activity and decomposition oforganic matter in the sediment. Addition of algaeenhanced the release of CO₂, NH⁺ ₄ ando-P. The excess inorganic C, N and P released inamended non-inhabited sediment after 36 dayscorresponded to 65, 31 and 58% of the C, N and P inthe added algae. In sediment inhabited by Chironomus plumosus the corresponding numbers were147, 45 and 73%, indicating that mineralization oforganic matter also from the indigenous sediment poolwas stimulated by chironomids. This revised version was published online in July 2006 with corrections to the Cover Date.     

The effect of tubificids and chironomids on particle redistribution of lake sediment

The effects of chironomid larvae,Chironomus plumosus, and tubificid worms,Limnodrilus spp., on particle redistribution in lake sediment were investigated experimentally using pots containing sediments obtained from Lake Suwa, Japan. The chironomids and tubificids increased the water content of surface sediment. The chironomid larvae had no effect on particle size distribution, while tubificids continuously accumulated small particles on the surface sediment through their selective feeding activity. Particles larger than 0.125 mm were buried at a sediment depth of 6 cm. In Lake Suwa, long diatom frustules, large plant debris and blue-green algal flocs were found to accumulate in the deeper layer of the lake sediment inhabited by tubificids at high density.     

Midge (Chironomidae,Chaoboridae, Ceratopogonidae) assemblages and their relationship with biological and physicochemical variables in shallow,polymictic lakes

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Factors influencing the species composition,distribution and abundance of benthic invertebrates in the profundal zone of a eutrophic northern lake

Factors influencing the species composition, distribution and abundance of benthic invertebrates were determined in a eutrophic subarctic lake from April 1978 to April 1979. Collections were made at five stations located at depths of 4 to 13 m. The largest populations of up to 5 × 10³ animals m^–2 were found in the deepest part of the lake. of the 24 species recorded in this area, the chironomidsProcladius denticulatus, Dicrotendipes modestus, Chironomus decorus andGlyptotendipes barbipes were most common. The strong development of benthos in the profundal zone was attributed to a consistently large supply of food and warm (4 °C) winter temperatures on bottom. Slightly smaller populations (up to 4 × 10³ animals, m^–2), composed of 19–23 species, occurred in shallower water, a reflection of lower (1.5 °C) winter temperatures. In the anoxic northern part of the lake, only 4–8 species were found in low numbers (400–1 000 animals m^–2). This was likely due to low (<5% saturation) oxygen levels in water and high organic content (18.5%) of the sediments.     

The effect of a predatory leech,Nephelopsis obscura,on mortality,growth, and production of chironomid larvae in a small pond

Summary The effect of a predatory leech, Nephelopsis obscura, on survivorship, growth, and production of chronomid larvae was studied by enclosure experiments carried out in a small pond. The prey population was composed almost entirely of the tubiculous, microphagous chironomid larvae, Chironomus riparius and Glyptotendipes paripes. Nephelopsis significantly reduced chironomid survivorship within the enclosures, and accounted for most of the measured mortality of fourth instar larvae. The cropping by Nephelopsis was not significantly biased toward either prey species. In long-term experiments (66 d) chironomid biomass in enclosures without leeches reached much higher levels than in enclosures containing Nephelopsis. This increase in biomass was due to growth of surviving larvae, rather than recruitment, since emergence and oviposition were not going on during the course of the experiments. The enhanced survivorship of larvae within leech-free enclosures was eventually accompanied by reduced growth and specific production (daily production/biomass) for C. riparius, which made up about 90% of the larval population. Growth and specific production of G. paripes (10% of larval population) was not affected. Short-term experiments (25 d) involving manipulation of densities and species ratio (9:1 CR:GP and 1:9 CR:GP) of larvae revealed that growth of the majority species was strongly influenced by larval density, whereas growth of the minority species was not. The same pattern was observed both in the presence and in the absence of Nephelopsis and was a result of differences in resource utilization between the two species. In the shortterm experiments, growth rates estimated for larvae exposed to leeches were significantly less than those for larvae in leech-free enclosures. This could be due either to size-biased consumption of larvae by Nephelopsis, or possibly a disturbance factor leading to reduced larval food intake and/or increased metabolic costs.     

Contribution to the knowledge of female west palaearctic Glyptotendipes Kieff. (Diptera, Nematocera, Chironomidae)

The characters of the genitalia of the females of Glyptotendipes glaucus (Meigen, 1818), Glyptotendipes pallens (Meigen, 1804), Glyptotendipes paripes (Edwards, 1929), Glyptotendipes gripekoveni (Kieffer, 1913), Glyptotendipes aequalis (Kieffer, 1922), Glyptotendipes barbipes (Staeger, 1839) and Glyptotendipes salinus Michailova, 1987 are described. A key for the identification of these species is given.The taxonomic status of Glyptotendipes aequalis is discussed.     

Relationship between δ13C of chironomid remains and methane flux in Swedish lakes

1. Methanogenic carbon can be incorporated by methane‐oxidising bacteria, leading to a ¹³C‐depleted stable carbon isotopic composition (δ¹³C) of chironomids that feed on these microorganisms. This has been shown for the chironomid tribe Chironomini, but very little information is available about the δ¹³C of other abundant chironomid groups and the relationship between chironomid δ¹³C and methane production in lakes. 2. Methane flux was measured at the water surface of seven lakes in Sweden. Furthermore, fluxes from the sediments to the water column were measured in transects in two of the lakes. Methane fluxes were then compared with δ¹³C of chitinous chironomid remains isolated from the lake surface sediments. Several different chironomid groups were examined (Chironomini, Orthocladiinae, Tanypodinae and Tanytarsini). 3. Remains of Orthocladiinae in the seven study lakes had the highest δ¹³C values (?31.3 to ?27.0‰), most likely reflecting δ¹³C of algae and other plant‐derived organic matter. Remains of Chironomini and Tanypodinae had lower δ¹³C values (?33.2 to ?27.6‰ and ?33.6 to ?28.0‰, respectively). A significant negative correlation was observed between methane fluxes at the lake surface and δ¹³C of Chironomini (r = ?0.90, P = 0.006). Methane release from the sediments was also negatively correlated with δ¹³C of Chironomini (r = ?0.67, P = 0.025) in the transect samples obtained from two of the lakes. The remains of other chironomid taxa were only weakly or not correlated with methane fluxes measured in our study lakes (P > 0.05). 4. Selective incorporation of methane‐derived carbon can explain the observed correlations between methane fluxes and δ¹³C values of Chironomini. Remains of this group might therefore have the potential to provide information about past changes in methane availability in lakes using sediment records. However, differences in productivity, algal δ¹³C composition and the importance of allochthonous organic matter input between the studied lakes may also have influenced Chironomini δ¹³C. More detailed studies with a higher number of analysed samples and detailed measurement of δ¹³C of different ecosystem components (e.g. methane, dissolved inorganic carbon) will be necessary to further resolve the relative contribution of different carbon sources to δ¹³C of chironomid remains.     

Distribution and seasonal abundance of benthic macroinvertebrates in a subtropical Florida lake

We studied the distribution and seasonal abundance of benthic macroinvertebrates from July 1975 through September 1976 in a hypereutrophic lake in subtropical Florida. The benthic community was comprised principally of oligochaetes (56.1%), chironomids (37.1%), and chaoborids (5.7%). Numbers of taxa and mean densities correlated negatively with depth and positively with mean grain size of the substratum and dissolved oxygen concentration at the mud-water interface. Seasonal abundances and life history information obtained for the predominant species of Chironomidae (Polypedilum halterale, Glyptotendipes paripes, Chironomus crassicaudatus, Cryptochironomus fulvus, C. blarina, Cladotanytarsus sp., Procladius culiciformis, and Coelotanypus concinnus) indicated that all of these species are multivoltine with rapid generation times. Larval lengths of life at summer temperatures, 27–31 °C, ranged from 14–22 days indicating that sampling in subtropical lakes should be at short intervals (approximately 3 days) if the life cycles, ecology, and function of the components of the benthic community is to be understood.     

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.     

Factors influencing the distribution of chironomids and oligochaetes in profundal areas of Spanish reservoirs

A canonical correlation analysis was performed in order to explain the distribution and abundance of the profundal macrozoobenthos in 71 Spanish reservoirs. Two sets of variables were considered: one set consisted of subgroups of benthic organisms based on their feeding strategy, and the other consisted of environmental variables related to the trophic state of the reservoir.The results showed that densities of phytophagous chironomids decreased with water depth and increased with oxygen content in the bottom waters, while Tanypodinae densities increased with rising densities of phytophagous chironomids and/or oligochaetes, revealing their strong dependence on the abundance of available prey. The abundance of oligochaetes and phytophagous chironomids showed a disjunct distribution with season and depth, suggesting the importance of food quality and sediment texture.Oxygen availability and water depth acted as governing factors for the zoobenthos. Chlorophyll-a was of secondary importance and correlated inversely with the abundance of benthic fauna, especially in summer. Autogenic production coupled with inputs of allochthonous coarse particulate organic matter and a long stratification period lead to long lasting anoxia that reduces the benthic fauna in Spanish reservoirs.Only up to 18% of the variability in the biological data could be associated with the environmental factors considered. Thus, factors other than the trophic state are much more important in reservoirs. Fluctuations of the water level and inputs of organic and inorganic materials coming from the basin are greater than in lakes and act as stressing factors for the benthic fauna.     

The life-history,distribution, and production of Chironomus riparius and Glyptotendipes paripes in a prairie pond

C. riparius and G. paripes exhibited univoltine life-cycles in Stephenson Pond; pupation, emergence and oviposition occurred mainly during May, and both species overwintered as mature fourth instar larvae. The marjority of larval growth for both species took place during the fourth instar stage (August–October), and growth and production were very low during late May to mid July when only young instars were present. Low production occurred during an interval when sestonic chlorophyll a concentration was very low, and the high production period corresponded to the Aphanizomenon bloom (August) and the autumn diatom pulse. None of the growth and production parameters investigated were correlated with temperature at the mud-water interface. Tube structure and behavior of the larvae indicate that G. paripes larvae are filter-feeders, whereas, C. riparius larvae are deposit-feeders.     

Chironomus species (Diptera: Chironomidae) in the profundal benthos of Spanish reservoirs and lakes: factors affecting distribution patterns

• 1 The taxonomy, distribution and habitat preferences of Chironomus larvae from profundal benthos of reservoirs and natural lakes in Spain are presented. The 114 water bodies studied were medium to large reservoirs and lakes throughout Spain, and were visited on at least two occasions (summer and winter). Out of the natural lakes investigated, Chironomus larvae were found only in one karstic lake (Banyoles), and to date, these have not been found in the high mountain lakes of the Spanish Pyrenees or Sierra Nevada, or in Spain's largest oligotrophic lake (Sanabria).
• 2 Multiple regression with environmental data from reservoirs showed that the density of Chironomus larvae was inversely correlated with water depth, temperature and sulphide content of the bottom water, but positively correlated with alkalinity and particulate organic nitrogen (PON) content of bottom water. Thus, the density of Chironomus larvae was higher in shallow reservoirs from the eastern part of Spain (calcareous water and higher PON), but markedly lower in southern reservoirs (higher temperature and higher sulphide).
• 3 According to morphological and cytological characteristics, three species of this genus have been identified in the samples examined: C. bernensis, C. plumosus and C. nuditarsis. Reasons for the absence of other widespread European species are discussed.
• 4 Using canonical correspondence analysis to relate species and environmental variables, C. bernensis showed a significant preference for reservoirs with higher oxygen concentrations in bottom water and lower total reactive phosphorus (TRP) content at the surface than those reservoirs where C. plumosus‐type were found. This suggests that the smaller C. bernensis prefers mesotrophic reservoirs and the larger C. plumosus‐type (C. plumosus + C. nuditarsis) prefers eutrophic reservoirs. Taking into account the TRP/depth index for those samples collected at depths below the thermocline, and thus, where oxygen can be a limiting factor, C. bernensis and C. nuditarsis were present in reservoirs in which the index was low (< 1:170 for C. bernensis and >1:800 for C. nuditarsis), while C. plumosus thrived in reservoirs with higher index values (<1:20). These differences may reflect the preference of C. bernensis and C. nuditarsis for detrital material, while C. plumosus prefers phytoplankton. These observations are in agreement with the depth distribution of each species: C. plumosus was restricted to reservoirs less than 46 m deep, whereas the other two species (i.e. C. bernensis and C. nuditarsis) were found up to depths of 65 and 80 m, respectively.
• 5 Data from a transect (5–20 m deep) in Lake Banyoles are consistent with the general pattern described in reservoirs. In this lake, a high density of C. bernensis was recorded from littoral and sublittoral areas with abundant oxygen (above or close to the thermocline), whereas C. plumosus was scarce in several parts of the lake, especially in oxygen‐deficient areas during the summer. This karstic, oligotrophic lake has a low TRP/depth index, and therefore, we expected C. bernensis to be dominant in the littoral, as is the case. Nevertheless, the severe depletion of oxygen caused by meromixis, and the high hydrogen sulphide concentration benefit C. plumosus in deeper zones. Chironomus nuditarsis was scarce in Lake Banyoles and only found at considerable depths.

     

Distribution and Diversity of Cyanobacteria and Eukaryotic Algae in Qinghai–Tibetan Lakes

Cyanobacteria and eukaryotic algae are important primary producers in a variety of environments, yet their distribution and response to environmental change in saline lakes are poorly understood. In this study, the community structure of cyanobacteria and eukaryotic algae in the water and surface sediments of six lakes and one river on the Qinghai–Tibetan Plateau were investigated with the 23S rRNA gene pyrosequencing approach. Our results showed that salinity was the major factor controlling the algal community composition in these aquatic water bodies and the community structures of water and surface sediment samples grouped according to salinity. In subsaline–mesosaline lakes (salinity: 0.5–50 g L^?1), Cyanobacteria (Cyanobium, Synechococcus) were highly abundant, while in hypersaline lakes (salinity: >50 g L^?1) eukaryotic algae including Chlorophyta (Chlorella, Dunaliella), Bacillariophyta (Fistulifera), Streptophyta (Chara), and Dinophyceae (Kryptoperidinium foliaceum) were the major members of the community. The relative abundance ratio of cyanobacteria to eukaryotic algae was significantly correlated with salinity. The algae detected in Qinghai–Tibetan lakes exhibited a broader salinity range than previously known, which may be a result of a gradual adaptation to the slow evolution of these lakes. In addition, the algal community structure was similar between water and surface sediment of the same lake, suggesting that sediment algal community was derived from water column.     

The influence of chironomus plumosus larvae on the exchange of dissolved substances between sediment and water

4th instar Chironomus plumosus larvae (about 1000·m^–2) were added to tubes containing sediment and overlying water. At a temperature of 20°C the larvae greatly increased the trasnport of silica, phosphorus and iron from the sediment to the water. Oxygen concentrations did not influence the exchange of silica. For two non-calcareous sediments the exchange of phosphorus and iron was much higher under anaerobic than under aerobic conditions while the difference was small for sediment from a hardwater lake. Exchange of inorganic nitrogen was little influenced by added chironomid larvae.     

Effects of resuspension and eutrophication level on summer phytoplankton dynamics in two hypertrophic areas of Lake Taihu,China

The composition of summer phytoplankton communities in two highly eutrophic areas of Lake Taihu, a large, shallow water body in China (area 2,338 km²; mean depth 2.0 m), was studied in relation to selected environmental variables. Both areas have multiple uses, including drinking water supply, fisheries, and recreation. The first area, Meiliang Bay, is much closer to central lake and subjected to wind-driven sediment resuspension, while the second, at Wuli Lake, is less affected by wave action but is considered more nutrient-rich due to its proximity to human inhabitation. By comparing two lake areas, we attempt to advance the understanding and management of different regions of shallow lakes, mainly differing in resuspension levels and eutrophication levels. A comparison of these two lake areas indicated a higher standing crop of phytoplankton in Wuli Lake than in Meiliang Bay. Canonic Correspondence Analysis indicated that nutrient status and sediment resuspension were the principal factors influencing phytoplankton composition in both areas. The dominant algal group in Meiliang Bay comprised immobile R-type strategists (Reynolds in Growth and reproductive strategies of freshwater phytoplankton, 1988), including filamentous diatoms and green algae (Aulacoseira granulate and Planctonema lauterbornii), and recruitment-beneficial RS-type strategists, mainly the colonial cyanobacteria (Microcystis spp.). The community of Wuli Lake was dominated by S-strategists, principally motile flagellates (Cryptomonas spp.) and C-strategists including small nonmotile centric diatoms and chlorococcal green algae.     

Benthos of Lake Mikri Prespa (North Greece)

An extensive sampling of Lake Mikri Prespa was carried out in summer 1990, aimed at evaluating benthic abundance, recording the species involved and detecting their response to some environmental variables. Three oligochaete species are endemic to the region,Potamothrix prespaensis, Psammoryctides ochridanus variabilis andPsammoryctides ochridanus typica. The bathymetric curve of the benthos resembles that of eutrophic lakes. A great abundance ofChironomus plumosus andChaoborus flavicans occurred. Four community patterns were revealed by TWINSPAN analysis, division level two, explicitly discriminated by the influence of substrate type, depth, C/N ratio and pH. Species richness and diversity was minimal in the profundal and maximal in the shallows, whilst the equitability index was high in all site groups. Depth and bottom type were the most important variables in defining species distribution. More precisely,Chironomus plumosus, Chaoborus flavicans andPotamothix hammoniensis were ubiquitous, preferring mostly deep waters.Tanytarsus sp.,Einfeldia dissidens, Cladopelma lateralis, Tanypus punctipennis andPotamothrix prespaensis favoured shallower and less productive waters with a bottom rich in decaying plant fragments, and thePsammoryctides species showed a littoral origin and regular occurrence on gravel beds. A few species displayed a particular distribution with changes of depth, C/N ratio and pH.     

USING EPILITHIC ALGAL COMMUNITIES TO ASSESS TROPHIC STATUS IN IRISH LAKES1

Relationships between taxonomic composition of shallow epilithic algal communities and nine environmental variables in 32 lakes of different trophic states in Ireland were explored using gradient analysis. A canonical correspondence analysis using four representative environmental variables, alkalinity (correlated with pH and conductivity), maximum phytoplankton chl a (CHL_max) (correlated with total P, total N, and chl), turbidity, and water color explained 21% of the variance in taxa distributions. The first two axes were significant and accounted for 77% of the variance in the periphyton–environmental relationship. The first axis was strongly related to alkalinity and color, which reflected geology and land use in the watersheds. The second axis was most correlated with CHL_max, and separation of lakes corresponded to their Organization for Economic Cooperation and Development (OECD) trophic classification based on water chemistry. Eutrophic lakes were characterized by cyanobacteria taxa and Stigeoclonium sp. Diatoms and desmids were generally more abundant in oligotrophic and mesotrophic lakes. Values for diatom trophic indices were poor indicators of trophic state. Weighted averaging regression and calibration techniques were used to develop transfer functions between 84 taxa and total P, total N, and CHL_max. The total P inference model predicted OECD trophic classification correctly for 84% of the lakes. Values for taxa preferences resulting from such models can provide the foundation for biomonitoring schemes using extant periphyton communities. The turnover time of periphyton taxa should integrate changes in environmental conditions at a temporal scale intermediate to surface‐sediment fossil diatom assemblages and water column variables, which may be more appropriate for detecting annual changes.     

Differences in feeding behaviour among Chironomus species revealed by measurements of sulphur stable isotopes and cadmium in larvae

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