Microdistribution of chironomids (Diptera: Chironomidae) in Alpine streams: an autoecological perspective

Chironomid communities from three glacial and three non-glacial high mountain streams in three Alpine river basins were analyzed (Conca, Niscli, Cornisello, NE Italy, 46°N, 10°E). Eighteen sampling reaches belonging to five stream types (kryal, subkryal, glacio-rhithral, kreno-rhithral, outlet) were investigated. At each reach, geomorphological, physical, chemical and biological data were collected. Field surveys were carried out during three periods per year from 1996 to 1998: immediately after spring snowmelt, in mid-summer and in early autumn. In all, 439 zoobenthos samples were collected from 5 to 10 microhabitats of 0.1m² in each reach and date using a standard pond net (mesh size of 250m). About 50% of individuals collected were chironomids (26673 specimens, 53 taxa), with densities ranging from 4 to 2652indm^–2. With few exceptions, they dominated as number of taxa and individuals in all reaches. Chironomid subfamilies Diamesinae and Orthocladiinae were most abundant, especially in glacial reaches, where Diamesa spp. constituted up to 100% of the total fauna. Chironomid distribution was analyzed in relation to 37 abiotic variables, referring to stream origin, hydrology, geomorphology, physics and chemistry. Diamesa steinboecki, D. latitarsis gr. A and Pseudokiefferiella parva were the taxa best associated with glacial conditions (i.e. high channel instability or presence of bedrock, high suspended solids and total phosphorous content, low conductivity and silica content, highly variable diel discharge and low mean temperature), while Pseudodiamesa branickii, Corynoneura spp., Eukiefferiella spp., Parorthocladius nudipennis, Tvetenia calvescens/bavarica, Thienemaniella spp. and Micropsectra atrofasciata were mostly associated with non-glacial conditions. Substratum particle size, water depth, current velocity, the presence of riffles/pools and of mosses/algae (Hydrurus foetidus) were the major factors affecting microdistribution of chironomids in the investigated streams.     

The chironomid fauna of four shallow,humic lakes and their representation by subfossil assemblages in the surficial sediments

A comparison of the present chironomid fauna of four shallow, humic lakes with the subfossils of surficial sediments is made. The chemistry and benthos of the lakes investigated varied greatly. Results indicate a reasonable correlation between biocenoses and thanatocenoses, but a marked under-representation ofProcladius is apparent. It is suggested that, at least in shallow, humic lakes,Procladius be excluded in attempts to reconstruct chironomid communities from subfossils.     

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.     

Ecological influences on larval chironomid communities in shallow lakes: implications for palaeolimnological interpretations

1. To correctly interpret chironomid faunas for palaeoenvironmental reconstruction, it is essential that we improve our understanding of the relative influence of ecosystem variables, biotic as well as physicochemical, on chironomid larvae. To address this, we analysed the surface sediments from 39 shallow lakes (29 Norfolk, U.K., 10 Denmark) for chironomid head capsules, and 70 chironomid taxa (including Chaoborus) were identified. 2. The shallow lakes were selected over large environmental gradients of aquatic macrophytes, total phosphorus (TP) and fish communities. Redundancy analysis (RDA) identified two significant variables that explained chironomid distribution: macrophyte species richness (P < 0.001) and TP (P < 0.005). Generalised linear models (GLM) identified specific taxa that had significant relationships with both these variables. Macrophyte percentage volume infested (PVI) and species richness were significant in classifying the lake types based on chironomid communities under twinspan analysis, although other factors, notably nutrient concentrations and fish communities, were also important, illustrating the complexities of classifying shallow lake ecosystems. Lakes with plant species richness >10 all had relatively diverse (Hill’s N2) chironomid assemblages, and lakes with Hill’s N2 >10 all had TP <250 μg L⁻¹ and total fish densities <2 fish per m². 3. Plant density (PVI), and perhaps more importantly species richness, were primary controls on the distribution of chironomid communities within these lakes. This clearly has implications for palaeoenvironmental reconstructions using zoobenthos remains (i.e. chironomids) and suggests that they could be used to track changes in benthic/pelagic production and could be used as indicators of changing macrophyte habitat. 4. Measuring key biological gradients, in addition to physicochemical gradients, allowed the major controls on chironomid distribution to be assessed more directly, in terms of plant substrate, food availability, competition and predation pressure, rather than implying indirect mechanisms through relationships with nutrients. Many of these variables, notably macrophyte abundance and species richness, are not routinely measured in such studies, despite their importance in determining zoobenthos in temperate shallow lakes. 5. When physical, chemical and ecological gradients are considered, as is often the case with palaeo‐reconstructions rather than training sets chosen to maximise one gradient, complex relationships exist, and attempting to reconstruct a single trophic variable quantitatively may not be appropriate or reliable.     

The underestimated role of temperature–oxygen relationship in large‐scale studies on size‐to‐temperature response

The observation that ectotherm size decreases with increasing temperature (temperature‐size rule; TSR) has been widely supported. This phenomenon intrigues researchers because neither its adaptive role nor the conditions under which it is realized are well defined. In light of recent theoretical and empirical studies, oxygen availability is an important candidate for understanding the adaptive role behind TSR. However, this hypothesis is still undervalued in TSR studies at the geographical level. We reanalyzed previously published data about the TSR pattern in diatoms sampled from Icelandic geothermal streams, which concluded that diatoms were an exception to the TSR. Our goal was to incorporate oxygen as a factor in the analysis and to examine whether this approach would change the results. Specifically, we expected that the strength of size response to cold temperatures would be different than the strength of response to hot temperatures, where the oxygen limitation is strongest. By conducting a regression analysis for size response at the community level, we found that diatoms from cold, well‐oxygenated streams showed no size‐to‐temperature response, those from intermediate temperature and oxygen conditions showed reverse TSR, and diatoms from warm, poorly oxygenated streams showed significant TSR. We also distinguished the roles of oxygen and nutrition in TSR. Oxygen is a driving factor, while nutrition is an important factor that should be controlled for. Our results show that if the geographical or global patterns of TSR are to be understood, oxygen should be included in the studies. This argument is important especially for predicting the size response of ectotherms facing climate warming.     

Environmental controls on the distribution and diversity of lentic Chironomidae (Insecta: Diptera) across an altitudinal gradient in tropical South America

To predict the response of aquatic ecosystems to future global climate change, data on the ecology and distribution of keystone groups in freshwater ecosystems are needed. In contrast to mid‐ and high‐latitude zones, such data are scarce across tropical South America (Neotropics). We present the distribution and diversity of chironomid species using surface sediments of 59 lakes from the Andes to the Amazon (0.1–17°S and 64–78°W) within the Neotropics. We assess the spatial variation in community assemblages and identify the key variables influencing the distributional patterns. The relationships between environmental variables (pH, conductivity, depth, and sediment organic content), climatic data, and chironomid assemblages were assessed using multivariate statistics (detrended correspondence analysis and canonical correspondence analysis). Climatic parameters (temperature and precipitation) were most significant in describing the variance in chironomid assemblages. Temperature and precipitation are both predicted to change under future climate change scenarios in the tropical Andes. Our findings suggest taxa of Orthocladiinae, which show a preference to cold high‐elevation oligotrophic lakes, will likely see range contraction under future anthropogenic‐induced climate change. Taxa abundant in areas of high precipitation, such as Micropsectra and Phaenopsectra, will likely become restricted to the inner tropical Andes, as the outer tropical Andes become drier. The sensitivity of chironomids to climate parameters makes them important bio‐indicators of regional climate change in the Neotropics. Furthermore, the distribution of chironomid taxa presented here is a vital first step toward providing urgently needed autecological data for interpreting fossil chironomid records of past ecological and climate change from the tropical Andes.     

When water interacts with temperature: Ecological and evolutionary implications of thermo‐hydroregulation in terrestrial ectotherms

The regulation of body temperature (thermoregulation) and of water balance (defined here as hydroregulation) are key processes underlying ecological and evolutionary responses to climate fluctuations in wild animal populations. In terrestrial (or semiterrestrial) ectotherms, thermoregulation and hydroregulation closely interact and combined temperature and water constraints should directly influence individual performances. Although comparative physiologists traditionally investigate jointly water and temperature regulation, the ecological and evolutionary implications of these coupled processes have so far mostly been studied independently. Here, we revisit the concept of thermo‐hydroregulation to address the functional integration of body temperature and water balance regulation in terrestrial ectotherms. We demonstrate how thermo‐hydroregulation provides a framework to investigate functional adaptations to joint environmental variation in temperature and water availability, and potential physiological and/or behavioral conflicts between thermoregulation and hydroregulation. We extend the classical cost–benefit model of thermoregulation in ectotherms to highlight the adaptive evolution of optimal thermo‐hydroregulation strategies. Critical gaps in the parameterization of this conceptual optimality model and guidelines for future empirical research are discussed. We show that studies of thermo‐hydroregulation refine our mechanistic understanding of physiological and behavioral plasticity, and of the fundamental niche of the species. This is illustrated with relevant and recent examples of space use and dispersal, resource‐based trade‐offs, and life‐history tactics in insects, amphibians, and nonavian reptiles.     

Distribution and production of the larval chironomid populations in Tjeukemeer

The distribution and production of larval chironomids in Tjeukemeer, a shallow, moderately large and wind exposed lake, were estimated between 1969 and 1972. The following seven species together contributed more than 95% of all the individuals collected: Chironomus anthracinus, Chironomus plumosus, Cladotanytarsus mancus, Glyptotendipes pallens, Polypedilum nubeculosum, Polypedilum scalaenum and Stictochironomus histrio. Density and biomass distribution depended generally on the substrate type and its stability. The standing crop biomass was highest in the reed-beds and in a narrow 100 m wide inshore zone and low in the large offshore zone. Wind induced wave action, causing substrate instability, is likely to be the main factor responsible for these low standing crops. The mean annual larval production in the lake was: 1969, 1.8; 1970, 0.4; 1971, 1.7 and 1972, 3.7 g dry wt m^–2.     

Identification of a biomanipulation‐driven regime shift in Lake Vesijärvi: implications for lake management

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Recovery from acidification in boreal lakes inferred from macroinvertebrates and subfossil chironomids

Acidification of waters and soils caused by emissions and the long-range transport of air pollutants has been a serious worldwide problem during the last decades. The extent of the acidification problem in Finnish acid-sensitive forest lakes was examined in the Acidification Research Project (HAPRO) in the mid-1980s. The recent decline in the emissions of air pollutants has resulted in the chemical recovery of watersheds in many regions, and the present work on the recovery processes in acidified Finnish headwater lakes (REPRO) was launched to examine whether the chemical recovery has already been accompanied by biological recovery. The patterns of recovery were studied by re-sampling littoral macrozoobenthos in a subset of the previously sampled HAPRO lakes. Paleolimnological samples were taken in order to assess the possible dependence of lacustrine chironomid communities on the changing degree of acidification. Acid sensitive and moderately acid sensitive benthic species revealed slight recovery in the formerly most acidic (pH 5.5) but recently recovered lakes. The most significant factors affecting the response of benthic communities were increased mean lake pH and decreased labile aluminium concentration. Paleolimnological chironomid analysis revealed a slight response along the pH gradient, but also significant structural similarity between the present and pristine chironomid assemblages. This implies that no major changes in chironomid communities of these acidic lakes have occurred during the past centuries. The alternative future trends and threats to biological recovery in small headwater lakes are discussed.     

Environmental implications of pollen spectra in bat droppings from southeastern Spain and potential for palaeoenvironmental reconstructions

Pollen was analysed from bat guano from nine caves in southeastern Spain and surface soils in their immediate surroundings. We compare the pollen spectra of 34 modern dung samples from the nine caves with one modern surface pollen sample from each cave. The contents suggest reasonable pollen diversity and richness, including anemophilous and zoophilous pollen types. Since the latter is usually under-represented in atmospheric pollen, the guano spectra therefore appear to reflect the vegetation more effectively than normal surface soil samples. Despite health hazards such as histoplasmosis, the difficulties of obtaining bat guano in deep caves and possible interpretational concerns relating to behaviour and feeding habits of different bat species, this material can be very useful in palaeoecological research provided that the dung was fossilized under favourable environmental conditions that allowed the preservation of pollen.     

Middle‐Late Ordovician iron‐rich nodules on Yangtze Platform,South China,and their palaeoenvironmental implications

Iron‐rich nodules occur on the Yangtze Platform, South China, especially along the marginal area, during the late Middle and early Late Ordovician (Darriwilian–Sandbian). Based on the study of samples from the Daling section, Anhui Province, these nodules include: (1) oncoids; (2) coated bioclasts; and (3) coated micritic intraclasts, all of which potentially have a similar biological origin. The nodule‐bearing strata were deposited in a midouter ramp setting, indicating sediment starvation and open marine conditions, possibly combined with anoxia in deeper basinal areas and cool water upwelling during the formation of the nodules, rather than terrestrial input or local tectonic movement. The widespread distribution of iron‐rich nodules at this time was probably favoured by high sea level and cooling climate, conditions that simultaneously favoured the evolutionary radiations of the GOBE in South China. Thus, these nodules form a time‐specific signature of a critical time in the evolution of the marine ecosystem in the context of local environmental background along the marginal Yangtze Platform.     

Different temperature perception in high‐elevation plants: new insight into phenological development and implications for climate change in the alpine tundra

In alpine habitats, predicted warmer and longer growing seasons will influence plant phenology, with important implications for species adaptation and vegetation dynamics. However, little is known on the temperature sensitivity of different phenophases and on the characteristics allowing phenological variation among and within species. By integrating interannual micro‐climatic variability with experimental warming, we explored how the phenology of three alpine species is influenced by temperature and what mechanisms underlie intra‐ and inter‐specific phenological differences. The present study demonstrated that alpine plants have different temperature responses during their reproductive cycle, do not have constant thermal thresholds and heat‐use efficiencies to achieve the seed dispersal stage and can change their temperature sensitivity to flower along snowmelt gradients. In addition, the length of the reproductive cycle, which proved to be species‐specific under experimental warming, does not seem to be the only life‐history trait under selective pressure due to the short‐length of the snow‐free period. In a warming climate scenario, the phenology of sexual reproduction will be considerably altered, and alpine plants may be subjected to changes in population dynamics driven by altered perception of environmental cues appropriate for coordinating the timing of key life‐history events.     

The case for studying tadpole autecology,with comments on strategies to study other small,fast-moving animals in nature

Two of the most fundamental questions in tadpole biology, also applicable to most small, under-studied organisms are: (1) ‘Why are they built the way they are?’ and (2) ‘Why do they live where they do?’ Regrettably, despite significant progress in most aspects of tadpole biology, the answers to these questions are not much better now than they were in the last century. We propose that an autecological approach, that is the careful observation of individuals and how they interact with the environment, is a potential path towards a fuller understanding of tadpole ecomorphology and evolution. We also discuss why more attention should be given to studying atypical tadpoles from atypical environments, such as torrential streams, water-filled cavities of terrestrial plants and wet rock surfaces neighbouring streams. Granted, tadpoles are rare in these settings, but in those unusual habitats the physical environments can be well described and characterized. In contrast, the more common ponds where tadpoles are found are typically too structurally complex to be easily delineated. This makes it difficult to know exactly what individual tadpoles are doing and what environmental parameters they are responding to. Our overall thesis is that to understand tadpoles we must see exactly what they are doing, where they are doing it, and how they are doing it. This takes work, but we suggest it is feasible and could greatly advance our understanding of how anuran larvae have evolved. The same strategies for studying tadpoles that we encourage here can be applied to the study of many other small and fast-moving animals.     

Phylogenetic diversity in freshwater‐dwelling Isochrysidales haptophytes with implications for alkenone production

Members of the order Isochrysidales are unique among haptophyte lineages in being the exclusive producers of alkenones, long‐chain ketones that are commonly used for paleotemperature reconstructions. Alkenone‐producing haptophytes are divided into three major groups based largely on molecular ecological data: Group I is found in freshwater lakes, Group II commonly occurs in brackish and coastal marine environments, and Group III consists of open ocean species. Each group has distinct alkenone distributions; however, only Groups II and III Isochrysidales currently have cultured representatives. The uncultured Group I Isochrysidales are distinguished geochemically by the presence of tri‐unsaturated alkenone isomers (C_37:3b Me, C_38:3b Et, C_38:3b Me, C_39:3b Et) present in water column and sediment samples, yet their genetic diversity, morphology, and environmental controls are largely unknown. Using small‐subunit (SSU) ribosomal RNA (rRNA) marker gene amplicon high‐throughput sequencing of environmental water column and sediment samples, we show that Group I is monophyletic with high phylogenetic diversity and contains a well‐supported clade separating the previously described “EV” clade from the “Greenland” clade. We infer the first partial large‐subunit (LSU) rRNA gene Group I sequence phylogeny, which uncovered additional well‐supported clades embedded within Group I. Relative to Group II, Group I revealed higher levels of genetic diversity despite conservation of alkenone signatures and a closer evolutionary relationship with Group III. In Group I, the presence of the tri‐unsaturated alkenone isomers appears to be conserved, which is not the case for Group II. This suggests differing environmental influences on Group I and II and perhaps uncovers evolutionary constraints on alkenone biosynthesis.     

Ultraviolet and temperature effects on planktonic rotifers and crustaceans in northern temperate lakes

1. Anthropogenic stressors such as climate change, ozone depletion and acidification may act in concert to alter ultraviolet (UV) light and temperature regimes in freshwater ecosystems. These physical and chemical changes will inevitably affect zooplankton community dynamics, but little is known about their relative effects on different species in natural communities. During spring, species that migrate to surface waters to take advantage of warmer temperatures may be especially vulnerable as UV levels can be high. 2. The objective of this study was to investigate the in situ effects of UV and temperature on a natural assemblage of planktonic rotifers and crustaceans during the spring. We performed in situ exposure experiments in two lakes with different surface temperatures. 3. Exposure to UV had a significant effect on the abundance and/or reproduction of four rotifers: Gastropus spp., Kellicottia bostonensis, Kellicottia longispina, Keratella spp.; two cladocerans: Holopedium gibberum, Daphnia catawba, and one copepod: Leptodiaptomus minutus. Incubation under cooler temperatures had a negative effect on K. longispina and H. gibberum. Temperature and UV had a significant interactive effect on abundance and/or reproduction of L. minutus and Ploesoma truncatum. Our results indicate that changes in underwater UV and temperature can significantly influence the composition of the zooplankton community and ultimately food web dynamics.     

Inferring past zooplanktivorous fish and macrophyte density in a shallow lake: application of a new regression tree model

1. Eutrophication has a profound effect on the biological structure and function of shallow lakes, altering the composition and abundance of submerged macrophyte and fish assemblages. Relatively little is known, however, about decadal to centennial‐scale change in these important aspects of shallow lake ecology. 2. Established palaeolimnological inference models are limited to reconstructing a single variable. As macrophyte and zooplanktivorous fish abundance exert dual and interacting controls on cladoceran assemblages a single variable inference model may contain significant error. To obviate this problem, we applied a new cladoceran‐based multivariate regression tree (MRT) model to cladoceran subfossil assemblages from dated cores from a small shallow lake (Felbrigg Lake, U.K.) to assess long‐term change in fish and submerged macrophyte abundance. Plant macrofossil, chironomid and mollusc subfossil assemblages were also analysed to track changes in biological structure and function and to evaluate the inferences of the MRT model. 3. Over the 200+ year period covered by the sediment cores, there was good agreement in the timing and nature of ecological change reflected by the plant macrofossil, mollusc, chironomid and cladoceran data. The sediment sequence was divided into three dated zones: c. 1797–1890, c. 1890–1954 and c. 1954–present. Prior to 1890 plant‐associated mollusc, cladoceran and chironomid assemblages indicated a species‐rich macrophyte community; a scenario confirmed by the plant macrofossil data. From c. 1890 to 1954 macrophyte‐associated species of all three invertebrate groups remained abundant but the proportion of pelagic cladocerans rose. Post‐1954 mollusc and chironomid assemblages changed to sediment associated detrital feeders and the proportion of pelagic cladoceran taxa increased further. 4. The cladoceran‐based MRT model indicated a long period of stability, c. 1790–1927, characterised by abundant submerged macrophytes and zooplanktivorous fish. From c. 1927 to 1980, the MRT model inferred a decline in zooplanktivorous fish density (ZF) but relative stability in August macrophyte abundance. From 1980 to 2000, an increase in zooplanktivorous fish was inferred tallying well with available data on the fish population (since the 1970s), which indicated extirpation of perch in the 1970s and a subsequent increase in the rudd population. The model inferred little change in August macrophyte abundance until post‐c. 1980 at which point it indicated a decline. The surface sediment assemblage was placed in MRT group A, where submerged plants are absent or very rare in late summer in good agreement with current conditions at the site. 5. The MRT model, applied here for the first time, appears to have successfully tracked changes in macrophyte abundance and ZF over the last 200 years at Felbrigg Lake. The inferences agreed with historical observations on the fish community and the supporting palaeolimnological data. Given that multiple structuring forces shape most biological communities, the application of a model capable of allowing for this represents a significant advance in palaeolimnology.     

Ecosystem response to earlier ice break‐up date: Climate‐driven changes to water temperature,lake‐habitat‐specific production,and trout habitat and resource use

Climate warming has yielded earlier ice break‐up dates in recent decades for lakes leading to water temperature increases, altered habitat, and both increases and decreases to ecosystem productivity. Within lakes, the effect of climate warming on secondary production in littoral and pelagic habitats remains unclear. The intersection of changing habitat productivity and warming water temperatures on salmonids is important for understanding how climate warming will impact mountain ecosystems. We develop and test a conceptual model that expresses how earlier ice break‐up dates influence within lake habitat production, water temperatures and the habitat utilized by, resources obtained and behavior of salmonids in a mountain lake. We measured zoobenthic and zooplankton production from the littoral and pelagic habitats, thermal conditions, and the habitat use, resource use, and fitness of Brook Trout (Salvelinus fontinalis). We show that earlier ice break‐up conditions created a "resource‐rich" littoral–benthic habitat with increases in zoobenthic production compared to the pelagic habitat which decreased in zooplankton production. Despite the increases in littoral–benthic food resources, trout did not utilize littoral habitat or zoobenthic resources due to longer durations of warm water temperatures in the littoral zone. In addition, 87% of their resources were supported by the pelagic habitat during periods with earlier ice break‐up when pelagic resources were least abundant. The decreased reliance on littoral–benthic resources during earlier ice break‐up caused reduced fitness (mean reduction of 12 g) to trout. Our data show that changes to ice break‐up drive multi‐directional results for resource production within lake habitats and increase the duration of warmer water temperatures in food‐rich littoral habitats. The increased duration of warmer littoral water temperatures reduces the use of energetically efficient habitats culminating in decreased trout fitness.