古温度定量重建的良好代用指标———湖泊沉积摇蚊化石记录研究进展

（双翅目）摇蚊科昆虫对环境因子的变化相当敏感，其水生幼虫的头囊能保存于湖相沉积物中，在国际古湖泊和古环境研究中常被用作一种环境替代指标。因为不同种属的摇蚊对环境因子的最适值和耐受范围有较大差异，所以利用湖相沉积物中的摇蚊化石组合面貌的变化可达到古环境的定量重建。在欧洲和北美进行的许多研究已证实摇蚊是古温度的有效指示器，重建夏季温度相对准确可靠，比其它气候变化代用指标有优势，很可能成为国际上古气温重建所使用的首要生物代用指标。截止目前，该领域研究在国内尚未开展，我国湖泊众多，利用摇蚊研究古温度记录潜力巨大。     

The dynamics of chironomid assemblages and vegetation during the Late Quaternary at Laguna Facil, Chonos Archipelago, southern Chile

We compare high-resolution pollen and chironomid records from the last 15,000 yr in Laguna Facil, southern Chile. Major vegetation and chironomid changes are recorded between ca 14,900 and 14,700 cal. yr BP. During the Lateglacial, changes in the chironomid stratigraphy lag behind changes in the pollen stratigraphy suggesting that the chironomids are responding to changes in the tree canopy or in soil chemistry brought about by vegetational development. At about 7200 cal. yr BP there is a change in the chironomid stratigraphy in advance of changes in the vegetation. This suggests that the response is to regional climatic change. The relatively close correlation of the chironomid and pollen stratigraphies with changes in charcoal concentrations also implicates the importance of fire and/or vulcanism in influencing the dynamics of forest and limnological systems. There is no clear evidence of cooling during the Younger Dryas chronozone in Laguna Facil.     

Anthropogenic versus climatic control in a high-resolution 1500-year chironomid stratigraphy from a southwestern Greenland lake

We performed a high-resolution study of chironomid assemblages in a sediment core retrieved from Lake Igaliku in southern Greenland. The well-dated core is located within the former Norse Eastern Settlement and covered the last 1500 yr. The comparison of chironomid stratigraphy (PCA axis scores) with instrumental temperature data, land use history and organic matter in the sediment over the last 140 yr suggested that the primary changes in chironomid fauna in 1988 ± 2 yr were driven by the shift to modern agriculture in the catchment. This unprecedented change in chironomid fauna was most likely triggered by a shift in in-lake processes. Within the instrumental period, subtle variations in the chironomid assemblages that occurred before 1988 ± 2 yr were significantly correlated with summer temperatures even in times of traditional extensive sheep farming in the catchment. The relevance of the chironomid-derived climate signal over the last 1500 yr was supported by its good concordance with previous studies in west Greenland and in the Arctic. The chironomid assemblage therefore appeared to be a valuable proxy for climate changes within the Norse colony area. Synchronous changes in Norse diet and chironomid-reconstructed climate give new insights into the interplay of Norse society with climate.     

The lacustrine sediment record of Oberwinkler Maar (Eifel, Germany): Chironomid and macro-remain-based inferences of environmental changes during Oxygen Isotope Stage 3

The lacustrine record of Oberwinkler Maar (Eifel, Germany) is the northernmost continuous record documenting the Weichselian Pleniglacial in central Europe – a period characterized by multiple abrupt climate oscillations known as the Dansgaard/Oeschger cycles. Here, the results of a high-resolution study of chironomid remains are presented, with a focus on the earlier part of Oxygen Isotope Stage (OIS) 3 (60–50 kyr BP) covering four stadial/interstadial cycles. During the stadials, the chironomid fauna of the former lake was dominated by many cold-stenothermic chironomid taxa, indicating a cold, oligotrophic lake. The concentrations of chironomid remains were lower during the interstadials, and featured a higher number of warm-indicating taxa. This could have been the result of a higher summer temperature at the study site, but also of bottom-water anoxia, an increase in trophic state or a combination of all these factors. During the stadial intervals, a taxon that is restricted to (sub-)arctic environments is present in our record, suggesting a change in the temperature regime rather than in-lake processes as the driving mechanism for the changes in the chironomid record. Although, consistently, there was a response of the lake ecosystem to climate changes, the amplitude of this response was not constant. This study provides unique and independent evidence of the impact of Dansgaard/Oeschger climate variability on the European continent during the earlier part of OIS-3.     

Interglacial Chironomidae (Diptera) from Thule, Northwest Greenland: matching modern analogues to fossil assemblages

An analysis of subfossil insect remains (Diptera, Chironomidae) from an interglacial site at Narsaarsuk near Thule Air Base, NW Greenland, was undertaken to complement our understanding of last interglacial environments in the Arctic by analogue matching to modern chironomid assemblages. The subfossil larval midge head capsules were well preserved and 82% of the chironomid remains were identified as eight different extant chironomid taxa. The assemblage was dominated by the lotic Diamesa (43.8%), a number of lentic taxa (Hydrobaenus, Psectrocladius, Cricotopus/Orthocladius) and a few semi-aquatic taxa (Smittia, Chaetocladius). A single black fly head capsule (Diptera, Simuliidae) was registered. The interglacial sample was compared to subfossil chironomid assemblages from 42 lakes in West Greenland, two glacier lakes (with and without river influence) and a quantitative zoobenthos study from Narsaq Elv. Similarity analysis, analogue matching and multidimensional scaling suggest a lotic, cold, glacier-fed interglacial palaeo-biotope. Quantitative temperature reconstruction was not possible owing to a high dissimilarity to modern lentic chironomid assemblages from West Greenland. However, the simple numerical methods convincingly managed to reflect an interglacial river and stream environment, which can be difficult to document from other palaeoecological data.     

Chironomid-based Lateglacial climatic reconstruction for southeast Scotland

This paper presents the first chironomid-based climatic reconstruction for the UK Lateglacial, obtained from a lake basin in southeast Scotland, Whitrig Bog. Comparison of chironomid, lithological, geochemical, and pollen data reveals a marked vegetation lag behind the warming following deglaciation; warm-water chironomid taxa (e.g. chironomus) had replaced cold-water chironomid taxa (e.g. Paracladius) prior to the arrival of shrubs and trees and when local soils were still poorly developed. In addition to clear evidence for the Younger Dryas Stadial, chironomid data also reveal two short-term cold episodes that punctuate the Lateglacial Interstadial, both of which are also reflected in pollen and lithological data as reversals from open birch scrub/woodland to open herbaceous tundra coinciding with inwash of minerogenic matter into the basin. © 1997 John Wiley & Sons, Ltd.     

Chironomids as indicators of the Holocene climatic and environmental history of two lakes in Northeast Greenland

Schmidt, S., Wagner, B., Heiri, O., Klug, M., Bennike, O. & Melles, M. 2010: Chironomids as indicators of the Holocene climatic and environmental history of two lakes in Northeast Greenland. Boreas, 10.1111/j.1502‐3885.2010.00173.x. ISSN 0300‐9483. Two Holocene sediment sequences from arctic lakes on Store Koldewey, an island in Northeast Greenland, were investigated for fossil chironomid assemblages. A total of 18 and 21 chironomid taxa were identified in 290‐ and 252‐cm‐long sediment sequences from Duck Lake and Hjort Lake, respectively. The chironomid assemblages were very similar in the two lakes. Canonical correspondence analysis (CCA) was used to compare fossil chironomid assemblages from Store Koldewey with chironomid assemblages and environmental conditions presently found in Canadian Arctic lakes and, hence, to infer environmental changes for Northeast Greenland. The first chironomids appeared at c. 9500 cal. a BP in Hjort Lake, and 500 years later in Duck Lake. Taxa typical for cold and nutrient‐poor arctic lakes dominated the earliest assemblages. Chironomid assemblages with taxa typical of higher summer air temperatures and lakes with higher nutrient availability occur between 8000 and 5000 cal. a BP. This period probably marks the regional Holocene thermal maximum, which is relatively late compared with some palaeoenvironmental records from East Greenland. One possible reason could be the location of Store Koldewey at the very outer coast, with local climatic conditions strongly influenced by the cold East Greenland Current. From around 5000 cal. a BP, chironomid assemblages in Duck Lake and Hjort Lake again became more typical of those presently found in Northeast Greenland, indicating relatively cold and nutrient‐poor conditions. This shift coincides with an increase of ice‐rafting debris off East Greenland and an intensification of the East Greenland Current.     

Recent Progress and Perspectives in Paleoenvironmental and Paleoclimatic Research Based on Chironomidae (Diptera)

Chironomidae (Diptera) becomes a rapid developing proxy in the international paleolimnology in the recent 20 years due to its short life cycle, strong ability to move and sensitive response to environmental change. The main progress of paleolimnological research based on chironomid can be summarized as the following four aspects: ①The establishment of a large number of sample training sets and transfer functions, as well as the comparison and integration of regional databases; ②More attention on the role of internal lacustrine factors in controlling of the chironomid population; ③Attention on finer time scales since the modern interglacial period; ④Improvement of the experimental technology and numerical methods, and constant development of new indicators. The domestic related research started late, but databases in the middle and lower reaches of the Yangtze river, the Tibetan Plateau, the Yunnan Plateau and the region of inner Mongolia and Xinjiang has been established so far, and also some researches on the internal lacustrine factors on chironomid population has been carried out; paleoenvironmental reconstruction is mainly concentrated in the northwest arid areas and the Yunnan Plateau, and the research in eastern region is primarily focused on reflecting the effects of recent state of human activities on the lakes. Paleoenvironment and paleoclimate researches based on chironomid urgently need to ①Carry out more individual ecology work to accurately understand the relationship between the species and environmental factors, combined with the investigation of a broader area; ②Further improve the efficiency of experiments by new technology to enhance the resolution of identification; ③Pay attention to the application of the advanced quantitative ecology methods and the novel tools such as isotopic analysis; ④Attach great importance to the role of chironomid in the definition of lake natural state and the ecological restoration.     

摇蚊亚化石记录的苏干湖近千年来盐度变化研究

利用摇蚊亚化石重建湖泊古盐度进而分析区域干湿变化已成为国际上古气候重建研究的热点之一。本文利用苏干湖SG03I孔的摇蚊亚化石组合变化分析了湖泊水体近千年来的盐度波动过程,沉积序列可划分为3个摇蚊组合带,通过高咸水属种Orthocladius/Cricotopus和Psectrocladius barbimanus,以及低咸水属种Psectrocladius sordidellus和Procladius的相对丰度变化实现盐度重建。结果表明,990～1450A.D.期间高咸水种处于绝对优势,湖泊盐度较高;1450～1850A.D.期间低咸水种显著增加,指示了小冰期期间较高的入湖水量与蒸发量之比;1850～2002A.D.期间低咸水种丰度回落,湖泊水体盐度逐渐增加至现代水平。SG03I孔岩芯摇蚊指标与其他代用指标的结果有很好的对应关系,摇蚊亚化石组合变化揭示的苏干湖区小冰期相对湿润的气候状况在干旱区其他记录中也有反映,证实了摇蚊亚化石分析在西北干旱区古环境研究中的有效性。     

The chironomid (Insecta: Diptera) and other fauna of aMyriophylum spicatum L. plant bed in the lower Hudson River

The benthic fauna of a small cove of the Hudson River containing the aquatic plantMyriophyllum spicatum L. was studied for a one-year period. The fauna was characteristic of oligohaline zones of United States east coast estuaries. Total abundance of invertebrates retained on a 0.12 mm mesh sieve averaged 124,631 organisms m^?2 (sediment and plant populations combined) and ranged up to 196,000 m^?2. During the May–August period, invertebrates living on the plants comprised 16–35% of the invertebrate fauna in the cove. Chironomid larvae were the most abundant organisms on plants and the third most abundant in the sediments. Two assemblages of chironomid species were recognized; one lived solely in the sediments, the other lived primarily on the plants.Chironomus decorus andTanytarsus sp. dominated the former group andCricotopus sylvestris the latter. The chironomidDicrotendipes modestus utilized both habitats. During the May–August period, chironomid biomass on the plants comprised approximately 50% of total chironomid biomass in the cove. The mean dry-weight biomass of chironomids in the cove (1.6 g m^?2) is estimated to be sixteen times greater than that of the fauna in the deeper areas of the river. Because chironomid larvae are eaten by fish and invertebrates, shallow water regions with their rich chironomid (and other fauna) may contribute importantly to the trophic dynamics of estuarine systems. *** DIRECT SUPPORT *** A01BY009 00004     

Chironomid grazing on benthic microalgae in a Louisiana salt marsh

Field experiments were conducted to examine spatial and temporal variation in chironomid (predominantlyTanypus clavatus) abundance, and their trophic relationship with benthic microalgae. High performance liquid chromatography (HPLC) analysis of chironomid gut pigments indicated that diatoms comprised the bulk of the microalgae ingested by chironomids.¹⁴C-feeding studies were used to obtain quantitative estimates of chironomid, copepod, ostracod, and nematode grazing on benthic microalgae. Daily consumption of standing microalgal biomass by chironomids ranged from 0.12% (January) to 125% (May), but was highly variable. There were no significant diel, temporal (over the scale of months), or spatial patterns in individual chironomid grazing rates. There was significant temporal variation in the proportion of microalgal biomass consumed by the total meiofaunal assemblage, and highest grazing impacts occurred in May, coincident with high abundances of chironomids, harpacticoid copepods, and ostracods. The grazing impact of chironomids was comparable to or greater than that of other known grazers of microalgae (copepods, ostracods). Functional-response experiments performed in the laboratory revealed that chironomid ingestion rates increased with increasing food availability over short (1 to 2 h) time scales. Field data did not indicate a functional response to food availability over longer (mo) time scales, possibly because of other environmental incluences. Gut residence time (determined using fluorescently labeled beads) changed with variable feeding rates, which were in turn a function of variable food availability. Chironomid larvae have the ability to consume significant fraction of the microphytobenthos in absolute terms, and relative to other meiofauna, indicating that they are an important component of the salt marsh food web.     

Chironomid-inferred temperature changes of the last century in anoxic Seebergsee,Switzerland: assessment of two calibration methods

Chironomids (non-biting midges) can provide accurate climate reconstructions from the Late Glacial to the present. Until now, anoxic lakes have been avoided for temperature reconstructions since chironomid assemblages are sensitive to changes in oxygen concentrations in the hypolimnion. However, anoxic lakes may have varved sediments, providing the possibility for near-annual climate reconstructions. Here, we tested the applicability of two calibration methods to reconstruct mean July air temperatures from chironomid assemblages preserved in the sediments of the anoxic Seebergsee located in the northern Swiss Alps: a calibration-in-space approach and a calibration-in-time approach. The calibration-in-space approach (i.e. chironomid assemblages from surficial lake sediments (0–1 cm) calibrated against meteorological data) provided accurate inferences (i.e. similar temperature changes as measured at the closest meteorological station, and at regional stations) in the Seebergsee stratigraphy until anoxia increased in the lake. With the increase of anoxia, the chironomid-inferred temperatures were generally colder than measured temperatures. A calibration-in-time approach (i.e. calibration of chironomid assemblages in a time series against instrumental data from the closest meteorological station) provided accurate reconstructions (i.e. similar to the regional records) for the past 100 years, including the time period of inferred anoxia. However, its applicability should be further tested on longer temporal scales.     

Diagenesis or dire genesis? The origin of columnar spar in tufa stromatolites of central Greece and the role of chironomid larvae

Two Greek Pleistocene tufa stromatolites were examined petrographically and with stable isotope geochemistry to determine whether calcite spar is of primary or diagenetic origin. The younger (ca 100 ka) tufa from Zemeno primarily is micritic, with primary columnar calcite spar restricted to areas immediately above chironomid larval tubes. This relationship suggests that chironomid larval feeding behaviour is responsible for the development of Zemeno tufa columnar calcite, probably involving biological substances smeared onto the tufa surface. Most micritic crystals are not suitably oriented to allow later post‐depositional growth resulting in columnar fabrics. The older (ca 1 Ma) predominantly sparry tufa from Nemea contains some chironomid tubes and organic cyanobacterial filaments preserved in crystal fans but also contains many fabrics found in primary speleothem spar. Columnar spar here is unlikely to be the result of post‐depositional crystal growth. A comparison of stable isotopic trends between the two tufa deposits suggests that both contain interpretable seasonal trends and implies little or minor post‐depositional alteration of either tufa. Consequently, there is no basis for the common assumption that sparry tufa fabrics must be of diagenetic origin.     

Chironomid‐inferred late‐glacial summer air temperatures from Lough Nadourcan,Co. Donegal,Ireland

Western Ireland, located adjacent to the North Atlantic, and with a strongly oceanic climate, is potentially sensitive to rapid and extreme climate change. We present the first high‐resolution chironomid‐inferred mean July temperature reconstruction for Ireland, spanning the late‐glacial and early Holocene (LGIT, 15–10 ka BP). The reconstruction suggests an initial rapid warming followed by a short cool phase early in the interstadial. During the interstadial there are oscillations in the inferred temperatures which may relate to Greenland Interstadial events GI‐1a–e. The temperature decrease into the stadial occurs in two stages. This two‐stage drop can also be seen in other late‐glacial chironomid‐inferred temperature records from the British Isles. A stepped rise in temperatures into the Holocene, consistent with present‐day temperatures in Donegal, is inferred. The results show strong similarities with previously published LGIT chironomid‐inferred temperature reconstructions, and with the NGRIP oxygen‐isotope curve, which indicates that the oscillations observed in the NGRIP record are of hemispherical significance. The results also highlight the influence of the North Atlantic on the Irish climate throughout the LGIT. Copyright © 2010 John Wiley & Sons, Ltd.     

The Late Devensian Lateglacial palaeoenvironmental record from Whitrig Bog, SE Scotland. 2. Chironomidae (Insecta: Diptera)

Changes in chironomid midge larval assemblages in Lateglacial (c. 13.5–10 ka yr BP) lake sediments from Whitrig Bog are used to infer climatic (temperature) change. The earliest sediments contain few, predominantly cold stenothermic taxa. This fauna is replaced by an assemblage dominated by thermophilic taxa, indicating rising temperatures. The relatively warm Interstadial is punctuated by at least two brief cold oscillations which are characterized by the return of certain cold-water taxa and the demise of some elements of the thermophilic fauna. The earlier of the two oscillations was apparently shorter and colder than the second. The highest Lateglacial Interstadial temperatures were attained either shortly before or after the first cold oscillation. This timing of the Lateglacial thermal maximum is apparently later than has been previously inferred from fossil beetle data. The Lateglacial Interstadial is terminated by the Loch Lomond (Young Dryas) Stadial, which is indicated in this monolith by an abrupt return of cold stenothermic Chironomidae and the virtual elimination of thermophilic taxa. Temperatures during the Younger Dryas appear to have been colder than during either of the previous minor cold oscillations. Climatic inferences from chironomid analysis broadly support and augment conclusions drawn from sediment chemistry and palynological evidence derived from the same monolith, although there is evidence that the vegetation and chironomid responses to early postglacial warming were out of phase.     

Subfossil Chironomidae (Insecta: Diptera) along a latitudinal gradient in Finland: development of a new temperature inference model

Numerical techniques were used to study chironomid distribution and abundance in lakes from a 1000 km transect in Finland, with special interest on the effect of local summer air temperatures on chironomid assemblages. The final aim of the study was to develop a chironomid‐based palaeotemperature inference model. The dataset consisted of 82 lakes (of which 77 were used in the model after deletion of outliers), with catchments spanning from boreal coniferous forests to mountain birch woodland and tundra vegetation. Numerical analysis showed that the mean July air temperature was the most significant variable explaining the distribution and abundance of chironomids in Finnish lakes. Weighted‐averaging partial least squares techniques were used to develop a palaeotemperature inference model for mean July air temperature reconstructions. The model performance statistics were favourable, with cross‐validated coefficient of determination (r²) of 0.78, root mean squared error of prediction of 0.721°C and maximum bias of 0.794°C. Based on these values, the transfer function is a valid means of performing quantitative palaeotemperature estimates in downcore studies. Copyright © 2008 John Wiley & Sons, Ltd.     

Invertebrate fossils (Insecta: Trichoptera,Diptera, Coleoptera) from the Pleistocene Scarborough Formation at Toronto,Ontario, and their paleoenvironmental significance

Larval caddisfly, chironomid, and beetle remains have been recovered from the Pleistocene Scarborough Formation in the Toronto region of southern Ontario. Three stratigraphic levels were sampled at the northeastern end of the Scarborough Bluffs; the youngest horizon yielded 16 chironomid taxa, 33 caddisfly taxa, and 28 beetle taxa, whereas the two older levels yielded somewhat less diverse assemblages. Only one taxon in each of the caddisfly and chironomid groups was identified from the presumed correlative beds at Woodbridge, Ontario, but numerous beetle fragments were recovered, several of which have been specifically identified and match species found previously in the upper part of the Scarborough Formation. The youngest sampled assemblage in the Scarborough Formation at the northeastern end of the Scarborough Bluffs is interpreted as indicating cool climatic conditions in a boreal forest environment, given the present-day distributions and feeding habits of these river, lake, and terrestrial taxa. The mean July temperature at this time was probably about 15°C, as compared to the present-day value of 20.5°C. The forest was poorer in deciduous species during deposition of the older part of the formation as preserved here. These results agree well with previous interpretations based on plant remains. We suggest that both aquatic and terrestrial insects are good indicators of macroclimate.     

Changes of treelines and alpine vegetation in relation to post‐glacial climate dynamics in northern Fennoscandia based on pollen and chironomid records

Palaeoclimatic records derived from a variety of independent proxies provide evidence of post‐glacial changes of temperature and soil moisture in northern Fennoscandia. We use pollen percentage, pollen influx, stomatal and chironomid records from Toskaljavri, a high‐altitude lake in northern Finland, to assess how treelines and alpine vegetation there have responded to these climate changes. The evidence suggests that the cool, moist climate of the early Holocene supported birch forest in the area 9600 cal. yr BP onwards and that a rise of temperature triggered the immigration of pine at 8300 cal. yr BP. At 6100–4000 cal. yr BP altitudinal treeline in the area was formed by pine, in contrast to the modern situation where mountain birch reaches a higher elevation. Alpine vegetation also demonstrates clear changes. Plant communities typical of dry, oligotrophic heaths of northern Fennoscandia expanded during the dry climatic period at 7000–4000 cal. yr BP and decreased in response to cooler and moister conditions after 4000 cal. yr BP. Alpine plant communities favouring moist sites show an inverse pattern, expanding after a change towards moister climate after 4000 cal. yr BP. In a redundancy analysis (RDA), a statistically significant proportion of the variability in the total chironomid assemblages was captured by changes in the pollen types reflecting alpine vegetation typical of moist sites. Although chironomid community changes appeared to follow the major patterns in the alpine vegetation succession, the present study does not support a direct link between the changing treeline position and chironomid stratigraphy. Rather, the data indicate that the terrestrial and aquatic environments have each responded directly to the same ultimate cause, namely changing Holocene climate. Copyright © 2002 John Wiley & Sons, Ltd.     

Lateglacial and early Holocene summer temperatures in the southern Swiss Alps reconstructed using fossil chironomids

We present a Lateglacial and early Holocene chironomid‐based July air temperature reconstruction from Foppe (1470 m a.s.l.) in the Swiss Southern Alps. Our analysis suggests that chironomid assemblages have responded to major and minor climatic fluctuations during the past 17 000 years, such as the Oldest Dryas, the Younger Dryas and the Bølling/Allerød events in the Lateglacial and the Preboreal Oscillation at the beginning of the Holocene. Quantitative July air temperature estimates were produced by applying a combined Norwegian and Swiss temperature inference model consisting of 274 lakes to the fossil chironomid assemblages. The Foppe record infers average July air temperatures of ca. 9.9 °C during the Oldest Dryas, 12.2 °C during most of the Bølling/Allerød and 11.1 °C for the Younger Dryas. Mean July air temperatures during the Preboreal were 14 °C. Major temperature changes were observed at the Oldest Dryas/Bølling (+2.7 °C), the Allerød/Younger Dryas (?2 °C) and the Younger Dryas/Holocene transitions (+3.9 °C). The temperature reconstruction also shows centennial‐scale coolings of ca. 0.8–1.4 °C, which may be synchronous with the Aegelsee (Greenland Interstadial 1d) and the Preboreal Oscillations. A comparison of our results with other palaeoclimate records suggests noticeable temperature gradients across the Alps during the Lateglacial and early Holocene. Copyright © 2011 John Wiley & Sons, Ltd.     

Wisconsinan Late-glacial environmental chage in New Brunswick: A regional synthesis

A synthesis is presented of the history of deglaciation, vegetation cover and climate in New Brunswick, Canada, during that part of the Wisconsinan Late-glacial period (12-9 ka BP) for which detailed records exist. Curves of inferred temperature variations in the region during this period are presented based upon qualitative interpretations of plant records and quantitative estimates based upon fossil chironomid assemblages.