Isolation of polymorphic microsatellite loci in three phantom midge species of the genus Chaoborus (Diptera: Chaoboridae)

Because of its widespread distribution in lakes and ponds Chaoborus is of great interest to many freshwater ecologists. Interestingly some species are restricted to small fish‐less water bodies, whereas other species live mostly in large lakes. To eventually test the genetic and evolutionary implications of these different lifestyles we identified microsatellite loci in three species in this preliminary study: C. obscuripes, C. crystallinus and C. flavicans. Using a biotin/streptavidin capture technique of repetitive sequences in a 96 well format, we obtained microsatellite‐enriched genomic libraries for all three species and identified six polymorphic microsatellite markers for each species.     

The Feeding of Chaoborus flavicans Meigen (Diptera,Chaoboridae). and its Predation on Lake Zooplankton

In a study of Chaoborus feeding in a eutrophic lake, selectivity was found to be positive with Crustacea (especially copepodit stages). and negative with Rotatoria. Daily food rations were about 20% for most of the feeding period, but higher (106%). during the month of intensive growth after hatching. Feeding intensity correlated positively with amount of food an temperature, and negatively with Chaoborus concentration. Elimination of Crustacea (in the epilimnion of the central zone of the lake). equalled about 30–40% of Crustacea production in June and September and slightly exeeded the August production (it was almost zero in the remaining months because Chaoborus larvae stayed at the bottom). This applies, however, only in the central zone – about 50% of the lake volume. Chaoborus probably influences both the density of zooplankton and the quantitative relations between zooplankton species.     

The seasonal dynamics and distribution of Chaoborus flavicans larvae in adjacent lake basins of different morphometry and degree of eutrophication

1.  Seasonal dynamics, spatial distribution and population size of the phantom midge Chaoborus flavicans in different parts of the eutrophic Lake Hiidenvesi (30.3 km²) were studied.
2.  Density of larvae was low in the shallow, most eutrophic parts of the lake, while the deep Kiihkelyksenselkä basin was inhabited by a dense population. In the deepest part of Kiihkelyksenselkä (33 m) density was 13 989 ± 3542 m^–2 in May, declined to 1102 ± 274 m^–2 in July and recovered to 7225 ± 1314 m^–2 by October. In spring and autumn the majority of larvae were benthic while, during high summer, few larvae were found in the sediment.
3.  Horizontal distribution fluctuated seasonally. On 3 June < 5% of the population inhabited areas shallower than 10 m. On 6 July the limnetic fraction was still restricted to regions deeper than 10 m, but 43% of benthic larvae were found between 6 and 10 m depths. In October both limnetic and benthic larvae were concentrated in areas deeper than 20 m.
4.  Within the lake, distribution was mainly regulated by stratification characteristics, degree of eutrophy being less important. The seasonal horizontal movements were probably induced by food shortage. Larvae could not meet their energetic demands in stratified areas and dispersed to shallower water, reducing predation risk by use of the benthic habitat.     

The dry weight estimate of chaoborus flavicans (Meigen) as a function of length and instars

Regression equations relating length to dry weight have been derived for the four larval instars of Chaoborus flavicans (Meigen). A general equation, in which all instars are lumped, was also developed, but produces biased results outside a rather narrow central interval.The rate of increase, in terms of weight, is minimal in first instar larvae and maximal in fourth instar larvae, while the second and third instars are both intermediate and very closely related to one another.Three critical points have been defined, corresponding to body lengths at which a particular instar has to molt in order to be able to continue growing optimally.

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Résumé Des équations de régression longueur-poids sec ont été établies pur les différents stades aquatiques de développement de Chaoborus flavicans (MEIGEN).La croissance pondérale, minimale pour les larves de premier stade, est maximale au cours du quatrième stade; les larves de deuxième et de troisième stade présentent des croissances comparables.L'emploi des équations propres à chaque stade est préférable à celui de l'équation moyenne établie pour l'ensemble des quatre stades larvaires.

     

Spatial and temporal distribution of benthic stages of Cyclops vicinus and Chaoborus flavicans in relation to abiotic factors and benthic fauna

The horizontal distributions of the benthic stages of Chaoborus flavicans and Cyclops vicinus were studied in a eutrophic stratified lake in the Massif-Central (France) over one year, at 5 stations from the shore to the centre of the lake. Their distribution was investigated in relation to temperature, dissolved oxygen, sediment grain-size and other benthic organisms. The dominant taxa of the benthic fauna of Lake Aydat were dipterans, crustaceans and oligochaetes and their distributions were independent of the grain size. In contrast to chironomids which preferentially inhabited the sublittoral zone, chaoborids and crustaceans were more numerous in the profundal zone. The sediment-dwelling oligochaetes remained numerous in both zones, according to the season. The fourth copepodite stages of Cyclops and Tubifex are tolerant to low oxygen concentrations in contrast to the fourth instar larvae of Chaoborus whose distribution was positively correlated with oxygen. The guts of these dipteran larvae were found to be empty and we assumed that, in contrast to the chironomids and oligochaetes, the resting stages of Cyclops vicinus and the benthic stages of Chaoborus flavicans did not use benthic resources. The former are activated at the autumn overturn, while the latter escape from the bottom at the start of the spring oxygen depletion. This suggests that physical factors are largely responsible for their reactivation. Both animals suffered of the effects of starvation and probably lost weight. The reactivation of the copepod at the autumn overturn would be facilitated by fluid mechanical disturbance. In addition, after the spring overturn, a small increase in temperature near 4 °C would be a reliable environmental signal for the dipteran. Food limitation does not occur, invertebrate predation pressure seems to be negligible and the predation by fish on the macrobenthic fauna and by chance on the meiofauna, clearly remains limited in both space and time. This revised version was published online in July 2006 with corrections to the Cover Date.     

Production ecology of phyto- and zooplankton in a eutrophic pond dominated byChaoborus flavicans (Diptera: Chaobolidae)

Primary production of phytoplankton and secondary production of a daphnid and a chaoborid were studied in a small eutrophic pond. The gross primary production of phytoplankton was 290 gC m⁻² per 9 months during April–December. Regression analysis showed that the gross primary production was related to the incident solar radiation and the chlorophylla concentration and not to either total phosphorus or total inorganic nitrogen concentration. The mean chlorophylla concentration (14.2 mg m⁻³), however, was about half the expected value upon phosphorus loading of this pond. The mean zooplankton biomass was 1.60 g dry weight m⁻², of whichDaphnia rosea and cyclopoid copepods amounted to 0.69 g dry weight m⁻² and 0.61 g dry weight m⁻², respectively. The production ofD. rosea was high during May–July and October and the level for the whole 9 months was 22.6 g dry weight m⁻².Chaoborus flavicans produced 10 complete and one incomplete cohorts per year. Two consecutive cohorts overlapped during the growing season. The maximum density, the mean biomass, and the production were 19,100 m⁻², 0.81 g dry weight m⁻², and 11.7 g dry weight m⁻²yr⁻¹, respectively. As no fish was present in this pond, the emerging biomass amounted to 69% of larval production. The production ofC. flavicans larvae was high in comparison with zooplankton production during August–September, when the larvae possibly fed not only on zooplankton but also algae.     

The Chaoborus (Diptera: Chaoboridae) component of the invertebrate drift of the Mwenda river,Lake Kariba,Central Africa

Chaoborus larvae were collected from the riverine drift of the Mwenda River during the 1972/3 rainy season. Large numbers of larvae were found in the early stages of flow following the 1972 dry season, fewer were found ten days later.Larvae which reached the lake in this way appeared to show no ill effects. Possible mortality arising prior to reaching the sampling point was doubtful since no dead larvae or parts were found in the material collected.     

Factors contributing to coexistence of Chaoborus flavicans and C. punctipennis (Diptera,Chaoboridae) in a small meromictic lake

In Crawford Lake, a small meromictic water body in southern Ontario, Canada, the life cycles of planktonic Chaoborus flavicans and C. punctipennis were out of phase by about two months, the former pupating from mid May through mid June and the latter from July through September. C. flavicans possibly produced a second annual generation. Fourth instars of both species were strong diel migrators and occupied similar strata at most times. C. punctipennis fourth instar diet consisted almost entirely of rotifers. C. flavicans ate rotifers but fed heavily also on daphnids in May and August. C. punctipennis fourth instars showed little growth until early spring, possibly owing to a sparseness of rotifers, then grew rapidly until pupation. C. flavicans had a slow, but relatively constant growth rate at all times during the open water season, presumably because its greater mouth gape allowed it a wider range of food items.     

Four different head shapes in Daphnia hyalina (Leydig) induced by the presence of larvae of Chaoborus flavicans (Meigen)

In many species of Daphnia spines, neck teeth, and enlarged or reshaped helmets are well-known as defences against invertebrate predators. Until now, Daphnia hyalina (Leydig, 1860), a common species in many European lakes, has appeared to be an exception to this rule.Here, we provide evidence that the larvae of Chaoborus flavicans (Meigen) also can induce morphological changes in D. hyalina. Specimens react morphologically to the presence of larvae in three ways: (a) by changes in the shape of head shield, (b) by the formation of spine(s) on the head, and (c) by the elongation of the tail spine. The frequency and intensity of these morphological changes are correlated positively with midge larvae densities. The most pronounced reactions occurred in young D. hyalina.The predator kairomone also induced changes in Daphnia body size.     

Ecological studies on Chaoborus (Diptera, Chaoboridae) in Lake George, Uganda

(1) Chaoborus anomalus and Chaoborus ceratopogones were found together throughout the year in Lake George, Uganda, the larvae forming 65-85% by number of the macro-benthos of the central lake area. (2) First and second instar larvae were wholly planktonic and third and fourth instar larvae mainly benthic during the day, migrating into the water column at night. A proportion of the third and fourth instar larvae were found in the water column during the day in the centre of the lake where light attenuation was greatest. C. anomalus showed a greater tendency to be benthic than C. ceratopogones. (3) Planktonic larvae were distributed in dense patches thought to be cohorts resulting from oviposition by swarms of adult females. The maximum density of planktonic larvae recorded was 31.7 × 10²/m² column. Third and fourth instar larvae were more evenly distributed in the benthos of the central lake area, the maximum density recorded was 79.9×10²/m². (4) First and second instar larvae occurred mainly in the inshore regions, very few were found in the centre of the lake. Oviposition is likely to take place inshore. The patches of planktonic larvae were not stationary. Dispersal of larvae away from the shore may be active or may result from complex oscillations of the water mass produced by storms. In C. ceratopogones the floating egg batches could he carried away from the north shore by currents produced by inflows and prevailing winds. (5) Adults of C. ceratopogones were more numerous during the rainy seasons than during the dry seasons. A similar, though less marked seasonality was found in the occurrence of C. anomalus adults. The biomass of benthic larvae of C. ceratopo-gones was estimated at 387.0 mg C/m² in the dry season, June 1970, and 130 0 mg C/m² in the rainy season, October 1970. That of C. anomalus was constant at 241.7 and 254.3 mg C/m² in June and October, 1970 respectively. (6) No difference was found in the horizontal distribution of the larvae, or in the size and weight attained by the various stages. (7) Interspecific differences were found in the mean number of eggs per adult female (309, C. anomalus and 224, C. ceratopogones); in the nature ofthe egg batches; n i the morphology of the larval mouthparts and pharyngeal sphincters, as well as i n the vertical distribution of the larvae and seasonal occurrence of the adults. On the basis of these differences the co-existing species are thought not to be competing for the same resources.     

Isolation of polymorphic microsatellite loci in the clear lake gnat and two other phantom midge species of the genus Chaoborus (Diptera: Chaoboridae)

Chaoborus is of great interest to many freshwater ecologists. The adults can become pests in certain areas in North America and the larvae are an important food source for fish. In this preliminary study, we identified variable microsatellite loci in three species: Chaoborus astictopus (H_E = 0.52–0.76), Chaoborus americanus (H_E = 0.46–0.80) and Chaoborus punctipennis (H_E = 0.66–0.81). Using a biotin/streptavidin capture technique of repetitive sequences in a 96‐well format, we obtained microsatellite‐enriched genomic libraries for all three species and identified six polymorphic microsatellite markers for each species. None of the primers did yield a polymerase chain reaction fragment in a cross‐species test.     

Zooplankton populations and the diets of three Chaoborus species (Diptera, Chaoboridae) in a tropical lake

SUMMARY. 1. Zooplankton populations in a small, natural, tropical lake are dominated by a few, small-sized taxa including the copepod Thermo- cyclops consimilis , the cladoceran Moina micrura and several rotifer species.
2. Moina micrura and adults of Thermocyclops consimilis undergo diel vertical migrations within the water column. Population densities of T. consimilis show marked intra-annual variations which may to some extent be related to variations in rainfall and to lunar periodicities In predator abundance.
3. The diets of Chaoborus lavae include other Chaoborus , Cladocera, Copepoda. Rotifera and the dinoflagellate Peridinium. The diets of late instars of the largest species, edulis , were dominated by Crustacea, while those of the two smaller species. C, ceratopogones and C. anomalus , were dominated by Rotifera and the dinoflagellate alga Peridinium , as were the diets of early instars of all species. Algae have not been previously reported to be a large component of the diet of Chaoborus populations in nature.
4. Some of the dietary differences among Chaoborus instars and species are related to the size of each prey species in relation to the mouth gape of each instar. However, there are also important differences in electivity among instars of different species of the same size.     

Persisting stemma neuropils in Chaoborus crystallinus (Diptera: Chaoboridae): Development and evolution of a bipartite visual system

Stemmata or “larval” eyes are of crucial importance for the understanding of the evolution and ontogeny of the hexapod's main visual organs, the compound eyes. Using classical neuroanatomical techniques, I showed that the persisting stemmata of Chaoborus imagos are connected to persisting stemma neuropils neighboring the first and second order neuropils of the compound eyes, and therefore also the imago possesses a stemma lamina and medulla closely associated with the architecture and the developmental pattern of those of the compound eyes. The findings are compared with other arthropods, e.g. accessory lateral eyes in Amandibulata and Myriapoda, suggesting some ancestral rather than derived character states. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Food selectivity and diel vertical distribution of Chaoborus edulis (Diptera, Chaoboridae) in Lake Malawi

1. Diel diet and vertical distribution patterns of the larval instars of Chaoborus edulis were studied in deep water near the central part of Lake Malawi, Africa.
2. First instar larvae contained very little food in their crops and probably depended on reserves from the egg. Second, third and fourth instars fed on zooplankton and were size-selective in their feeding. The mean size of prey eaten by the three instars was significantly different from each other, with larger instars feeding on larger prey. Smallest available prey was selected against and the upper size of prey was probably constrained by larval gape. Nauplii were not found in any of several thousand larvae examined. Phytoplankton did not form a significant part of the diet.
3. There was a progressive and related increase in diel periodicity in feeding and vertical migrations of successive instar stages. Fourth instars migrated particularly large distances. Such migrations removed them from their zooplankton food supply but avoided predators. A refuge from predators is probably found in or near the permanent zero oxygen boundary, at depths greater than 200 m.     

The estimation of crop evacuation rates in Chaoborus larvae (Diptera: Chaoboridae) using natural prey

SUMMARY. 1. Crop evacuation rates were estimated for the first time in Chaoborus larvae, using natural prey.
2. Fourth instar C. americanus Matheson digested copepods (Diaptomus leptopus S.A. Forbes) 48.7% faster than daphnids (Daphnia rosea Leydig) of similar size; meal size did not significantly affect the instantaneous rate of digestion (IRD) within each prey type. Prey specific IRD has not been reported before for zooplankton.
3. Prey specific differences in IRD require the use of natural prey when digestion experiments are to be used to estimate natural rates of food consumption of animals.     

Effects of competitors and Chaoborus predation on the cladocerans of a eutrophic lake: an enclosure study

The role of large laboratory grown food competitors ofthe genus Daphnia as well as the predationimpact of Chaoborus on the cladoceran communityof an eutrophic lake was assessed in five insitu enclosure experiments. The hypothesis tested wasthat the outcome of competition and gape-limitedpredation on cladocerans is size dependent. Accordingto the generally accepted assumptions on competitionand invertebrate predation, large-bodied cladocerantaxa were expected to be less affected by competingcongeners and by Chaoborus than weresmall-bodied taxa. Effects of the predator upon anassemblage of differently sized cladoceran taxa weremuch more pronounced than effects of competition.There was a tendency of predation and competitionimpact to decrease with cladoceran size, but predationpressure was also low for some small cladocerans andhigh for some large cladocerans. The general trendswere further obscured by factors not or indirectlylinked to body size.     

Larva-adult relationships in an ancestral dipteran: A re-examination of sensillar pathways across the antenna and leg anlagen of Chaoborus crystallinus (DeGeer, 1776; Chaoboridae)

 In one of his classical studies on insect metamorphosis, Weismann compared the imaginal anlagen of the ancestral phantom midge, Chaoborus, with those of advanced brachycerans. We have expanded his findings on the relationships between larval and imaginal organs using electron microscopy and cobalt backfilling of the antenna and leg anlagen and the axonal trajectories of corresponding larval sensilla. We show that both primordia are confluent with the larval antennae and ”leg” sensilla (an ancestral Keilin organ), respectively. These fully developed larval organs represent the distal tips of the imaginal anlagen rather than separate cell clusters. The axons of the larval antenna and leg sensilla project across the corresponding anlagen to their target neuromeres within the central nervous system (CNS). Within the discs, nerves composed of these larval axons, developing afferent fibres and efferences ascending from the CNS are found. Both the structure of the primordia and the axonal trajectories thus relate the situation found in advanced brachycerans with that seen in more ancestral insects. In addition, the larval antennae, legs, wings and even the eyes possess very similar afferent pioneer trajectories supporting the idea that the described pattern is generally used in the ontogeny of sensory systems. Received: 30 June 1998 / Accepted: 27 September 1998     

Primary producer dynamics associated with evaporative concentration in a shallow,equatorial soda lake (Lake Elmenteita,Kenya)

Lake Elmenteita (0°27S, 36°15E) lies on the floor of the rift valley at 1776 m above sea level in Kenya. As a consequence of lower than average rainfall, the mean depth decreased from 1.1 to 0.65 m during the study period (February 1973 to August 1974). The initiation of major biological changes coincided with a period of rapid evaporative concentration in 1973 (February to April) when the conductivity increased from 19.1 to 27.0 mmhmos cm^-1. Spirulina platensis, Spirulina laxissima and Anabaenopsis arnoldii decreased in abundance precipitously in parallel with large declines in chlorophyll a concentration and phytoplankton photosynthetic rates. Once the overall abundance of phytoplankton had declined and the transparency had increased, primary productivity by benthic algae increased significantly. Paradiaptomus africanus, the only copepod living in the lake, was abundant in February and March 1973, but was gone by May. Eight hypotheses to explain these changes are evaluated and converge on the suggestion that a rate of change of salinity greater than 5 mmhmos cm^-1 per month and a salinity exceeding 25 mmhmos cm^-1 cannot be tolerated by P. africanus and adversely effects the nitrogen fixer, A. arnoldii. Furthermore, the loss of P. africanus and oxygenation of the sediments by benthic algae reduce the rate of recyling of nutrients which alters phytoplankton abundance and species composition.     

Remarks on the morphology and biology of Cleigastra apicalis (Meigen, 1826) (Diptera, Scathophagidae)

The egg, second‐ and third‐instar larvae and puparium of Cleigastra apicalis are described for the first time. All pre‐imaginal stages are found on stems of the common reed affected by flies of the genera Lipara and Platycephala and the butterfly Arenostola phragmitidis. The larvae feed on dead plant and animal tissue and the excreta of other insects that live inside the stems of the common reed. Exceptionally they will scrape living plant tissue. The pupa is the overwintering stage.     

Subtle variation in phosphorus availability influences mating biology in Hyalella (Amphipoda: Hyalellidae) amphipods

Eutrophication caused by anthropogenic activity has had tremendous impacts on aquatic ecosystems across the world. A primary cause of this eutrophication is increased phosphorus run‐off into water bodies, resulting in alterations to the quantity and quality of food available for consumers. Previous studies have shown that large differences in the availability of phosphorus affect the expression of traits and mating biology at the individual level. In the present study, we manipulated the supply of phosphate, which altered the quantity and quality of food (e.g. periphyton) available to consumers, and documented the population‐level responses of Hyalella amphipods. We also performed a mating experiment to examine the effects of the availability of phosphorus during development on the expression of sexual traits, as well as phosphorus availability during mating on mating behaviour. The results obtained indicate that subtle variation in the availability of phosphorus has no effect on population density or sex ratio. The amount of phosphorus available during development and phosphorus available during mating, however, did affect mating behaviour. Furthermore, the patterns in mating behaviour observed were not attributable to phosphorus‐induced changes in sexually selected traits. The present study indicates that subtle changes in the availability of phosphorus can affect mating biology, possibly as a result of physiological or behavioural changes, although such effects may not necessarily be mediated by differential expression of sexually selected traits. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 878–888.