Refugia and habitat partitioning among midges (Diptera: Chironomidae) in rain-pools

ABSTRACT.

• 1 Rain-pools on granite exposures are common in tropical Africa and they are inhabited by the larvae of two midge species, Chironomus imicola Kieffer and Chironomus pulcher Wiedemann.
• 2 These pools differ from other ephemeral island habitats such as carrion, fallen fruit, sap flows, mushrooms and dung in their high degree of spatial predictability. Most pools are predictably inhabited by either C.imicola or C.pulcher. This finding is considered in the light of contemporary competition theory.
• 3 An arrangement whereby the same species repeatedly invades the same temporary habitat indicates that there is something about the habitat itself causing the discrimination. I suggest that degree of exposure to sunlight may provide a cue allowing females to discriminate between pools as oviposition sites.
• 4 Both species survive the long dry season by retreating to pools of river water left by receding rivers. Dispersal to rain-pools by emerging adults occurs during the following season with imicola occupying rain-pools remote from shaded rivers. Appropriately, measured against pulcher, the biology of imicola is that of a colonizing species.
• 5 The possibility is discussed that the chironomids in rock-pools do not survive to breed. Instead, these populations may be maintained externally, for example by invasions from breeding populations in the Great Lakes of Africa, where larval densities may be too low to be detected by sampling. The evolutionary implications of such a situation are considered.

     

Production and Ecology of Benthic Chironomid Larvae (Diptera) in Lake Hayes,New Zealand,a Warm-monomictic Eutrophic Lake

The distribution and abundance of larval chironomids in Lake Hayes were studied from December 1973 to March 1975. The mean annual production of the two dominant species, Chironomus zealandicus and Chironomus sp. a, was 29.2 g m⁻² dry weight which is approximately 4.3 % of the average annual phytoplankton production in the lake. A high annual P/B ratio of 18.5 is consistent with the multivoltine life cycle of C. zealandicus. Larval chironomid production in the second summer when Anabaena blooms were absent was only one quarter of that in the first summer and is consistent with the hypothesis that the production of benthic chironomids in Lake Hayes is closely linked to that of the phytoplankton through the sedimentation of autochthonous organic matter.     

Subsidies for production of Chironomus calligraphus larvae (Chironomidae,Diptera) in the laboratory

The objective of this study was to evaluate the effect of temperature, type of mineral substrate, and type and amount of feed on the growth of Chironomus calligraphus as a subsidy for the production of Chironomus larvae to feed the fish. Egg production was evaluated at 22, 25, and 28?°C on fine (0.074–0.210?mm), or silty sand (0.002–0.250?mm), and no mineral substrate as control. For larval production, 0.10, 0.20, and 0.30?mg/larvae of fish feed, poultry feed, or turkey manure were used as food. The highest number of egg masses was produced when mineral substrates and 25?°C were used, but the higher number of eggs/mass was registered in the control. Therefore, the total number of eggs and hatched larvae produced was the same in the presence or in the absence of mineral substrates (p?>?0.05). The best results for survival and growth were recorded for larvae reared with fish or poultry food. The increase in the amount of feed was correlated with weight gain only for larvae fed with poultry feed, with no reduction in survival.     

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.     

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.     

Ecological background and importance of the change of chironomid fauna (Diptera: Chironomidae) in shallow Lake Balaton

The objectives of this research were to record the changes in composition of the open-water, bottom-dwelling chironomid fauna in Lake Balaton between 1978–1984, to examine the causes of these changes, and to discover their significance in the life of the lake.The spatio-temporal dispersion of larvae is compared with the water and sediment quality of each basin in the lake. It is established that, under present conditions, nutrient status can be regarded as the chief environmental factor.Studies of population dynamics show that chironomids play a highly important role in preserving sediment quality. Chironomids are an essential element in the organic matter circulation of the lake. They dominate a sub-system that retards water quality degradation, and thus they play a prominent role in the natural prevention of eutrophication.     

Associations of Chironomidae (Diptera) of shallow,acid, humic lakes and bog pools in Atlantic Canada,and a comparison with an earlier paleoecological investigation

The composition of benthos in a series of 29 lake and peat pool sites is examined especially in relation to a successional gradient. The results indicate a sharp distinction between the fauna of peat pools and lakes. Chironomus, Psectrocladius, Monopsectrocladius, and Zalutschia are characteristic of bog lakes at the latter stages of their evolution and peat pools. Procladius and Tanytarsus dominate in most lakes. Between weakly acidic and strongly acidic lakes a sharp boundary exists for many other components of the benthos. Chaoborus occurs in strongly acidic lakes. Amphipoda, and Ephemeroptera are limited to weakly acidic or circum-neutral waters.A comparison of the results of this investigation with a parallel paleoecological study is made.     

Chironomid flies of the genus Glyptotendipes Kieffer (Diptera: Chironomidae) from India with a short account of their biology

Three new species, Glyptotendipes (Glyptotendipes) fumilatus, G. (G.) ingenius and G. (G.) validens are described. A short account of habitat, construction of dwellings, food and feeding of the larva and emergence of the adult has also been appended. A new dichotomous key to the Indian species is provided.     

The evolutionary history of two species of Orthocladiinae (Diptera: Chironomidae) from Lake Baikal (Eastern Siberia)

The evolutionary history of two species belonging to the genus Orthocladius van der Wulp, 1874 van der Wulp, F.M. (1874), ‘Dipterologische Aanteekneningen’, Tijdschrift voor Entomologie, 17, 109–148. [Google Scholar] (Diptera: Chironomidae) from Lake Baikal was investigated using the mitochondrial gene coding the first subunit of the cytochrome c oxidase (CO1 mtDNA). The phylogenetic analysis indicated that the Baikal Orthocladius species were divided into two well-defined clades where O. (Orthocladius) gregarius Linevitsh, 1970 was a sister species to Palaearctic O. (Orthocladius) nitidoscutellatus Lundstrom, 1915 and the O. (Eudactylocladius) sp. was a sister species to Nearctic O. (Eudactylocladius) subletteorum Cranston, 1998 Сranston, P.S. (1998), ‘Nearctic Orthocladius Subgenus Eudactylocladius Revised (Diptera: Chironomidae)’, Journal of the Kansas Entomological Society, 71(3), 272–295.[Web of Science ®] , [Google Scholar]. Divergence time estimates indicated that these species had been evolving independently for about 18 Ma (Neogene, Early Miocene), while emergence of the most recent common ancestors of the modern O. (Orthocladius) gregarius and O. (Eudactylocladius) sp. was dated to about 3.5 Ma (Neogene, Pliocene). The evolution of Baikal orthoclads occurred from the rheophilic fauna under conditions of global climate change during the geological history of the Baikal Depression in the Tertiary Period.     

Records of chironomids of the tribe Pentaneurini (Diptera: Chironomidae) in the Eastern Himalayas of India

Larva, pupa and female imago of Paramerina ampliseta n. sp. and Rheopelopia tuberculata (Chaudhuri & Debnath 1987), pupa and male imago of Paramerina clara n. sp. and pupa of Ablabesmyia (Ablabesmyia) alba Chaudhuri, Debnath & Nandi 1983 and Paramerina inficia Chaudhuri & Debnath 1985 are described from the Eastern Himalayas of India.     

Mariambera,a new genus of Orthocladiinae from Brazil (Insecta: Diptera,Chironomidae)

The genus Mariambera is erected based on the males of M. amazonica n. sp. from Amazonas and M. mariae n. sp. from Minas Gerais. The new genus is easily recognized by the distinctive anal point which has a rounded dorsal lobe with microtrichia and a triangular ventral lobe with very strong, marginal setae, in combination with bare eyes and wing membrane, squama with setae, antenna without strong apical seta, and apparently few, short, very weak, hair-like acrostichals in anterior 1/3 of scutum.     

The flightless marine midge Pontomyia (Diptera: Chironomidae): ecology,distribution, and molecular phylogeny

Pontomyia Edwards, 1926 (Diptera: Chironomidae) is a genus of exclusively marine flightless midges. There are four described species from the Indo‐Pacific, and one undescribed species known only from females, pupal skins, and larvae from the Atlantic/Caribbean. They are poorly known owing to their small size (～1.0 mm), extremely short adult life (< 3 h), and unusual habitat for an insect (coastal lagoons, bays, or rock pools). We reviewed scattered literature on their biology and systematics, presented photomicrographs of the male hypogium, and updated the geographic distribution of each species. We carried out the first molecular study to elucidate relationships among and within three of the species. Results from our four‐gene phylogenetic reconstruction using combined gene tree and species tree approaches showed that Pontomyia natans, Pontomyia oceana, and Pontomyia pacifica are each well‐supported clades, with P. natans as sister to P. oceana + P. pacifica. Genetic distances based on mitochondrial cytochrome oxidase I are extraordinarily large within P. natans and P. pacifica, which suggests that they may be cryptic species complexes. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 443–456.     

Distribution and production of chironomids (Diptera: Chironomidae) in a shallow, alkaline lake (Neusiedler See, Austria)

From July 1990 to July 1991 the benthic community of the open water zone of Neusiedler See, one of the largest shallow lakes in central Europe, was studied with special reference to the chironomids. Only 16 spp. of chironomids inhabited the sediment of the open water zone. The numerically dominant species were Tanypus punctipennis, Procladius cf. choreus, Microchironomus tener and Cladotanytarsus gr. mancus. Most invertebrates showed a distinct horizontal distribution. Species richness and abundance were highest on muddy and organically rich substrates near the reed belt. Chironomid densities in this area reached 54,000 ind m^–2 and biomass was 2.0 g dw m^–2. The two tanypod species accounted for more than 90% of the standing stock of the macrozoobenthos near the reed belt. The sediment of the open lake and of the eastern part of Neusiedler See was composed of compact clay and sand as a result of the erosion of fine material due to strong waves and currents. Individual densities in these areas were much lower. Production of the numerically dominant species T. punctipennis was estimated using the increment-summation method, whereas production of the remaining species was estimated using an empirically derived multiple regression. Mean annual production of chironomids exceeded 6 g dw m^–2 yr^–1 near the reed belt, but it reached only 0.55 g dw m^–2 yr^–1 in the open lake. These values are rather low compared with other lakes and can be explained by unfavourable sediment conditions due to wave action and by physiological stress due to the water chemistry.     

The distribution of Heterotrissocladius oliveri Saether (Diptera: Chironomidae) in Lake Michigan

Fifty one chironomid species were identified from 504 samples collected at depths ranging 8 to 267 m in Lake Michigan, U.S.A. Heterotrissocladius oliveri Saether occurred in 32% of these samples and had an average abundance of 22 m^–2 which was similar to other estimates from the Great Lakes. Maximum average lake-wide density was at 30 to 60 m (41 m^–2). At depths 60 m, H. oliveri was the dominant chironomid species comprising 75% of total Chironomidae. The substrate preference of H. oliveri differed within each depth regime considered: at 30–60 m, 2–3 ; at 60–120 m, 3–5 , 7–9 ; and at 120–180 m, 6–8 . Abundance was notably reduced at all depths in substrates characterized as medium silt (5–6 ). On a lake-wide basis, the distribution pattern suggested H. oliveri was most numerous from 30 to 60 m along the southwestern, eastern, and northern shorelines and at 60–120 m depths along the southern and eastern shorelines. Increased abundance in the South Basin was concurrent with evidence of increased sedimentation at 60 to 100 m. However, in several other areas of the lake, high densities were associated with medium to very fine sands relatively free of silts and clays. This observation suggested occurrence of H. oliveri was minimally affected by sediment type.Widely variable, but generally elevated water temperatures likely prevent H. oliveri from establishing a substantial population density at depths < 30 m. With increased depth, temperature fluctuation is negligible and food is more stable, though the source is variable. Factors limiting abundance of H. oliveri at depths 30 m were related to decreased food supply due to distance from shore, food sources of lower value (clays), and, most importantly, to reproductive replenishment.Although still oligotrophic in nature, high density occurrences in both high and low sedimentation areas of the lake suggest the trophic indicator status of H. oliveri might be broader than previously thought.     

Habitat preferences as patch selection of larval and emerging chironomids (Diptera) in a gravel brook

Four common species of larval and emerging chironomids were investigated during 1981 and between 1984/1985 in an alpine gravel brook. The emergence patterns ofN. dubius, O. rivulorum andH. ornaticollis were significantly affected by fluctuations of the water level, while rises in water temperature tended to trigger the peaks in emergence of the same species and ofO. frigidus. N. dubius exhibited its main emergence activity over areas of bed sediment which had a lower mean water velocity or were discontinuously flooded. On the other hand, the imagines ofO. frigidus, O. rivulorum andH. ornaticollis reached peak densities in zones of the main current channel in the Seebach. The larvae of these species inhabited horizontally and vertically different areas of bed sediment depending on the developmental stage. Early instars ofN. dubius andH. ornaticollis preferred zones near the banks and reached their maximum densities in sediment depths of 10 to 20 cm and 20 to 30 cm, respectively. In contrast, the larvae ofO. frigidus andO. rivulorum reached their highest densities in the first 10 cm of the sediments with significant preferences for the zones of stronger water flow.     

Laboratory investigation on community composition, emergence patterns and biomass of wood-inhabiting Chironomidae (Diptera) from a sandy lowland stream in Central Europe (Germany)

Wood-inhabiting chironomid communities were investigated from June 1998 to July 1999 by laboratory rearing of randomly collected submerged branches from a headwater and mid-reach site of a sandy lowland stream, separated by an impoundment used for fishery purposes. Total annual emergence (males and females) from headwater samples was higher (2551 ind m^–2 y^–1) compared to the mid-reach (1576 ind m^–2 y^–1), which could be due to disturbances caused by frequent impoundment openings resulting in high discharge events. Chironomid community from branches comprised three subfamilies, with Orthocladiinae (18 species, 2189 ind m^–2 y^–1) clearly predominating at the headwater (total of 36 species). Mid-reach samples (total of 48 species) showed similar emergent numbers of Orthocladiinae (19 species, 786 ind m^–2 y^–1) and Chironominae (26 species, 764 ind m^–2 y^–1). Tanypodinae were caught very rarely in the laboratory emergence (12 ind m^–2 y^–1 at both sites). Shannon-Wiener diversity index for the mid-reach chironomid community was higher (2.52) than for the headwater community (1.68). Chironomid species composition on woody debris was similar between stream sites, with a Sørensen index of 0.75, but showed different dominance structures indicated by a Wainstein index of 0.26. Total adult biomass (in the manner of dry mass) during the study period was higher for wood-dwelling chironomids from the headwater (158.2 mg m^–2 y^–1) compared to the mid-reach (123.8 mg m^–2 y^–1), but individual biomass was higher for mid-reach chironomids (0.079 mg ind^–1 vs. 0.062 mg ind^–1 at the headwater), indicating the predominance of larger species. Total biomass of wood-inhabiting chironomids in the investigated lowland stream was low compared to other field emergence studies, which could be attributed to the laboratory approach for investigating the emergence from a single substrate type (submerged wood) instead of the integrative field surveys where chironomids from all habitats were caught. Main reason for the lack of chironomid species closely associated to wood in this sandy lowland stream could be infrequent but episodic disurbances caused by the anthropogenic induced highly fluctuating discharge regime of the downstream study site.     

Laboratory investigation on community composition, emergence patterns and biomass of wood-inhabiting Chironomidae (Diptera) from a sandy lowland stream in Central Europe (Germany)

Wood-inhabiting chironomid communities were investigated from June 1998 to July 1999 by laboratory rearing of randomly collected submerged branches from a headwater and mid-reach site of a sandy lowland stream, separated by an impoundment used for fishery purposes. Total annual emergence (males and females) from headwater samples was higher (2551 ind m^–2 y^–1) compared to the mid-reach (1576 ind m^–2 y^–1), which could be due to disturbances caused by frequent impoundment openings resulting in high discharge events. Chironomid community from branches comprised three subfamilies, with Orthocladiinae (18 species, 2189 ind m^–2 y^–1) clearly predominating at the headwater (total of 36 species). Mid-reach samples (total of 48 species) showed similar emergent numbers of Orthocladiinae (19 species, 786 ind m^–2 y^–1) and Chironominae (26 species, 764 ind m^–2 y^–1). Tanypodinae were caught very rarely in the laboratory emergence (12 ind m^–2 y^–1 at both sites). Shannon-Wiener diversity index for the mid-reach chironomid community was higher (2.52) than for the headwater community (1.68). Chironomid species composition on woody debris was similar between stream sites, with a Sørensen index of 0.75, but showed different dominance structures indicated by a Wainstein index of 0.26. Total adult biomass (in the manner of dry mass) during the study period was higher for wood-dwelling chironomids from the headwater (158.2 mg m^–2 y^–1) compared to the mid-reach (123.8 mg m^–2 y^–1), but individual biomass was higher for mid-reach chironomids (0.079 mg ind^–1 vs. 0.062 mg ind^–1 at the headwater), indicating the predominance of larger species. Total biomass of wood-inhabiting chironomids in the investigated lowland stream was low compared to other field emergence studies, which could be attributed to the laboratory approach for investigating the emergence from a single substrate type (submerged wood) instead of the integrative field surveys where chironomids from all habitats were caught. Main reason for the lack of chironomid species closely associated to wood in this sandy lowland stream could be infrequent but episodic disurbances caused by the anthropogenic induced highly fluctuating discharge regime of the downstream study site.     

The karyotype of Axarus festivus (Say) and a comparison with species of Lipiniella (Chironomidae: Diptera)

Larvae of Axarus andLipiniella share several apparantly apomorphic features of the mouth-part structure, however adults ofLipiniella show possible relationships toDemeijerea andChironomus. In order to assist in refining the relationships ofAxarus toLipiniella the karyotype ofAxarus festivus was determined and compared to the karyotypes ofLipiniella arenicola andL. moderata. Larvae ofA. festivus from a population in Kansas were monomorphous, with 2n=8, the Ist, IInd and IIIrd chromosomes metacentric, and IVth acrocentric.Axarus festivus therefore differs fromL. arenicola in chromosome number (2n=6), however homologous sections of all chromosomes were identified. Inversions were detected in the Ist, IInd and IIIrd chromosomes ofA. festivus relative toL. arenicola. It was determined that both species have high functional activity, as indicated by the presence of three Balbiani rings, and more than one nucleolus per genome. Differing degrees of polyteny, a feature previously described forL. arenicola, were observed in the salivary glands, with highest degrees of polyteny in cells near the salivary duct. These similarities of chromosome structure indicate close genetic relationships betweenA. festivus andL. arenicola. However, we did not find evidence for similarity ofA. festivus toL. moderata, which supports the previous conclusions byKiknadze et al. (1989) regardingL. arenicola andL. moderata.     

Titimbera,a new genus of Orthocladiinae from South and Central America (Diptera: Chironomidae)

Titimbera n. gen. is erected based on the males of three new species from South and Central America: T. amazonica n. sp. from the Amazon region, Brazil; T. titi n. sp. from Venezuela and T. laselvensis n. sp. from Costa Rica. The combination of bare eyes and wing membrane; antenna without strong apical seta; scalpellate acrostichals in mid scutum; costa strongly extended; R₄₊₅ ending opposite to M₃₊₄; Cu₁ strongly curved to slightly sinuous; anal point sitting high on tergite IX, nearly parallel-sided with bluntly rounded apex; and club-shaped to subtriangular gonostylus with distinct heel will separate the genus from all other orthoclads.