Differential use of food resources by the instars of Chaoborus punctipennis

SUMMARY. 1. Differential use of food resources by instars I-1V of Chaoborus punctipennis was examined in a mesotrophic New Hampshire lake during midsummer when all instars coexisted. Diet composition and prey preference were quantified first, at night when all instars were feeding at the same depth, and second, during the day when the early instars (I-II) and instar III were segregated by depth. Relative abundance, per cent biomass, and per cent frequency of occurrence of soft and hard-bodied rotifers, flagellated phytoplankton, protozoans, and crustaceans in C. punctipennis diets were quantified using crop content analyses. 2. All four instars ingested large, flagellated phytoplankton (mainly Dinobryon and Ceratium). This unexpected result suggests that the effects of phytoplankton on Chaoborus growth and survival should be investigated. All instars also consumed rotifers (mainly Kellicottia, Gastropus, Polyarthra), but only instars III and IV fed upon crustaceans (mainly Daphnia, Bosmina and Diaphanosoma). Small rotifers (Gastropus spp., Keratella cochlearis, Trichocerca similis) occurred more frequently and were more abundant in early instar diets than late instar (III and IV) diets, whereas large rotifers (Asplanchna priodonta and Keratella crassa) were eaten only by instars III and IV. Zooplankton with gelatinous sheaths (e.g. Ascomorpha, Collotheca and Holopedium) were rarely ingested. 3. Developmental increase in gape diameter of C. punctipennis seemed to be the major proximate mechanism causing dietary differences among instars. Body widths of hard-bodied prey in C. punctipennis crops were always less than gape diameter. The relationship between prey body width and Chaoborus gape diameter, coupled with knowledge of prey escape behaviour, should be useful for predicting the presence of hard-bodied prey taxa in diets of other Chaoborus species. 4. All four instars of C. punctipennis selected soft-bodied, or weakly loricate, rotifer prey over crustaceans and phytoplankton. Early instars preferred the small rotifer T. similis and the protozoan Difflugia sp. to other rotifers and phytoplankton when feeding in the epilimnion or thermocline during the day or night. Late instars positively selected the rotifer, Asplanchna. Prey value (prey weight ingested per unit handling     

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

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

A comparison between predation effects on zooplankton communities by Neomysis and Chaoborus

Community level effects of predation by two invertebrate predators, the opossum shrimp (Neomysis intermedia), and the larva of the phantom midge (Chaoborus flavicans) were studied and compared. N. intermedia appeared abundantly in the shallow eutrophic Lake Kasumigaura and had a significant impact on the zooplankton community. The predation pressure by Neomysis was highest on cladocerans, followed by rotifers, and finally copepods. At high densities (maximum nearly 20 000 individuals m^–2), Neomysis excluded almost all cladocerans, rotifers and copepods from the lake.Zooplankton communities were established in experimental ponds, into which C. flavicans was introduced. The predator's density was around 1 individual l^–1, and was probably controled by cannibalism. Although Chaoborus larvae feed on various zooplankton species, their predation impact on zooplankton populations was markedly selective. They eliminated medium- and small-sized cladocerans and calanoid copepods from the ponds, but rotifers increased.Although the feeding selectivities of Neomysis and Chaoborus individuals were similar, the predation effects on zooplankton communities by the two predators were different.     

Prey utilization by Chaoborus punctipennis Say in a small,eutrophic reservoir

Prey selection by 2nd-, 3rd-, and 4th-instar larvae of Chaoborus punctipennis was evaluated by comparing larval crop contents to the species structure of the zooplankton community in a small, eutrophic lake. The rotifers, Brachionus caudatus and Trichocerca similis, were positively selected by and comprised most of the diet of 2nd- and 3rd-instar larvae. Rotifers comprised 67% of all prey items identified in the crops of even 4th-instar larvae. Whereas 2nd-and 3rd-instar larvae selected the two rotifers with equalfrequency, 4th-instar larvae selected Trichocerca significantly more frequently than they did Brachionus. Although copepods comprised only 8% of all prey identified in 3rd-instar crops, they comprised 33% of the crop contents of 4th-instar larvae. Both 3rd and 4th instars exhibited a positive electivity for Cyclops vernalis significantly more frequently than for Diaptomus pallidus. We conclude that the larvae of Chaoborus punctipennis can, and do, select between Trichocerca and Brachionus and between Cyclops and Diaptomus.The paper by Brooks and Dodson (1965) triggered a flurry of studies on aquatic predators and their ability to discriminate between available prey species. Although the original work dealt with vertebrate predators, it has become increasingly apparent that invertebrate predators may also exhibit considerable selectivity in choosing between prey categories. The larval stages of the various species of Chaoborus are ubiquitous lentic predators, especially in mesotrophic or eutrophic systems. Although there have been a number of recent publications which deal with the food habits and preferences of several species of Chaoborus, only a few studies have included analyses of larval crop contents and none of these have included Chaoborus punctipennis. This species is of particular interest because it is one of the smaller species in the genus, and perhaps for this reason, because it is frequently present in lakes which support large populations of planktivorous fish (von Ende 1979). The purpose of our research was to characterize the food habits and preferences of Chaoborus punctipennis in a small, eutrophic reservoir which supports a rather depauperate number of species of zooplankters.     

Photoresponses of late instar <Emphasis Type="Italic">Chaoborus punctipennis</Emphasis> larvae to UVR

With a few clear exceptions (e.g., Daphnia) it is uncertain if most aquatic invertebrates can detect and respond to ultraviolet radiation (UVR). It is known that many aquatic invertebrates are vulnerable to UVR and that anthropogenically-induced increases in surface UVR have occurred in recent decades. We examined the photoresponses of late larval instars of Chaoborus punctipennis to different combinations of UVA (320–400 nm), UVB (300–320 nm) and visible light (400–700 nm) to determine whether the larvae can detect and/or avoid UVR. To accomplish this, we exposed late instar C. punctipennis larvae to a directional light source of UVR only (peak wavelength at 360 nm), visible light only or visible plus various wavebands of UVR. We examined negative phototaxis for 10 min at a quantum flux of 2.62 x 10¹³ quanta s^–1 cm^–2 (S.D. = 3.63 x 10¹² quanta s^–1 cm^–2). In the dark, larvae stayed close to the surface of the experimental vessels. Under all treatments containing visible light the larvae exhibited negative phototaxis and occupied the bottom of the vessels. Under UVR only, the larvae occupied the middle of the water column. Our results suggest that late instar C. punctipennis larvae are unable to detect and avoid UVB and short UVA wavelengths but they can detect long UVA wavelengths.     

Environmental correlates of the intrinsic rate of natural increase in primates

Morphological and life history traits of two clones of the cladoceran Daphnia pulex were measured in the presence and absence of size-selective insect predators, the midge larva Chaoborus flavicans, which preys on small Daphnia, and the water bug Notonecta glauca, which preys on large Daphnia. The aim was to detect predator-induced phenotypic changes, particularly the effect of simultaneous exposure to both types of predators. Other work has shown that in the presence of Chaoborus americanus, Daphnia pulex produce a socalled neck spine which may carry several teeth. The morphological modifications are supposed to serve as an anti-predator device. Furthermore, females exposed to Chaoborus often delay their maturation; this has been interpreted as a cost that balances the benefits of the neck teeth. In this investigation, females of both clones produced fewer but larger offspring than control animals when reared in the presence of Chaoborus flavicans. The offspring showed the typical neck spine and delayed first reproduction. In the presence of Notonecta glauca, one of the clones produced more and smaller offspring, and maturation occurred at earlier instars. The other clone also produced more offspring than the control but there was no size difference. When both predators were present, in most cases the reactions of the daphnids were similar to those in the Notonecta experiment. The response to Chaoborus appeared to be suppressed. The observed modifications are interpreted as evolved strategies that reduce the impact of size-selective predation. They are consistent with predictions of life-history theory.     

Vertical migration of Chaoborus flavicans in a Scottish loch

The pattern of diel vertical migration of Chaoborus flavicans larvae in a shallow Scottish loch varies according to the instar and, in third and fourth instars, according to the season. The planktonic phase of the migration is not exclusively concerned with predation, the larvae feeding at least equally actively in the benthos. Although there is no conclusive evidence as to the role of vertical migration in Chaoborus flavicans, it has some of the characteristics of an epideictic display. Upward and downward locomotion can be induced experimentally in a plankton wheel by manipulating light intensity alone. However, the responses are complex and there is a great deal of individual variation, suggesting that additional factors are involved.     

Chaoborus predation and zooplankton community structure in a rotifer-dominated lake

During summer, Chaoborus punctipennis larval densities in the water column of fishless, eutrophic Triangle Lake become very high, and coincidently, the spined loricate rotifer Kelfcottia bostoniensis becomes the dominant zooplankter. Research was done to test the hypothesis that selective predation by Chaoborus on soft-bodied rotifers controls species dominance in the mid-summer zooplankton of this lake. In situ predation experiments showed positive selection by Chaoborus for the soft-bodied Synchaeta oblonga, negative selection for K. bostoniensis, and intermediate selection for Polyarthra vulgaris, a species with rapid escape tactics. However, during a 21 day in situ mesocosm experiment, zooplankton dominance and succession in Chaoborus-free enclosures was identical to that in enclosures with Chaoborus at lake density. Despite the selective predation, Chaoborus larvae may not exert significant top-down control on rotifers, whose intense reproductive output during mid-summer in temperate eutrophic lakes results in new individuals at rates that exceed predatory losses.     

The extant Chaoborus assemblage can be assessed using subfossil mandibles

1. Assemblages of Chaoborus were examined in 80 thermally stratified southern central Canadian Shield lakes to explore whether subfossil mandibles could be useful in assemblage‐level studies of Chaoborus. 2. Chaoborus (Sayomyia) (probably Chaoborus punctipennis in this study region) and Chaoborus flavicans were the most common taxa recorded, while Chaoborus trivittatus was rarer. Chaoborus americanus was not recorded in subfossil assemblages, because no fishless lakes were included in this study. Chaoborus flavicans had higher relative abundances (%) in lakes with higher dissolved organic carbon (DOC), probably because of reduced fish predation in less transparent water. 3. Results from logistic regression indicate that patterns of presence/absence for Chaoborus mandibles in the study lakes were influenced primarily by hypolimnetic oxygen concentration, probably because of the presence or extent of a hypolimnetic refugium from fish predation. 4. Chaoborus species richness in lakes, derived from subfossil assemblages, did not differ significantly from species richness estimates derived from plankton sampling with a net. Patterns of dominance and coexistence [e.g. the widespread co‐occurrence of C. flavicans and C. (Sayomyia)] determined from subfossil assemblages agreed with previous studies of the contemporary living assemblage. 5. These results suggest that subfossil assemblages may be used as an alternative to nocturnal plankton sampling to carry out research on the community ecology of Chaoborus. 6. We propose a hierarchical conceptual model of assemblage‐level patterns of Chaoborus in temperate lakes. Chaoborus americanus dominates in fishless lakes, whereas in lakes with fish Chaoborus is typically absent where there is no anoxic hyplimnion. In lakes with anoxic strata, C. trivittatus tends to dominate in lakes with few fish; in the remaining lakes, C. flavicans and C. (Sayomyia) dominate, although C. flavicans is more relatively abundant in lakes with lower water clarity (higher DOC).     

Zooplankton of a flooded opencast sulphur mine

The zooplankton community of a flooded opencast sulphur mine was investigated during two years. The complicated physical and chemical relationships in this water-filled pit caused an atypical plankton assemblage. At the time of the investigation 20 species of rotifers, 6 of Cladocera, and 8 species of Copepoda were found. A single Ostracod and Chaoborus flavicans were also noted in the samples. Usually the epilimnion was monopolised by rotifers whereas the hypolimnion had a more diversified assemblage of rotifers, cladocerans and copepods. The result of clustering analysis indicated that each predator was associated with its own group of prey. Discrimination analyses showed weak seasonal differences in the zooplankton and three zones in the vertical profile – epilimnetic, upper hypolimnetic and anaerobic. Principal component analysis reduced the chemical parameters to the main contributors of conductivity (the main anions and cations) plus toxic sulphide and hydrogen sulphide. The animals were classified by the first two components into two groups: cold stenotherms and medium-sized filtrators, such as Bosmina longirostris. A specific analysis of vertical profiles generated five variants of diel vertical migrations. When time–space distributions of the plankters were analysed the highest numbers of different species were usually in different times and depths. The epilimnion was inhabited by small fishes and was dominated by rotifers but was free from C. flavicans which inhabited the cold hypolimnion, together with the remaining invertebrate predators (copepods and Asplanchna).Chaoborus and the other invertebrate predators participated more evenly in the contributions of species to the total density. In the presence of invertebrate predators the structure of the rotifer population shifted from soft-bodied species to spined or fast-moving species. When the predation pressure was too strong, as in the hypolimnion, the soft-bodied species were, in practice, absent. When the pit was in an oligotrophic state the hypolimnion population was composed of up to 60–90% rotifers. When the trophic state shifted to mesotrophy, the proportion of cladocerans increased.     

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.

     

The biology of Brachycentrus subnubilus Curtis (Trichoptera) in the River Frome,Dorset

Aspects of life-history, oviposition and microdistribution within beds of Ranunculus penicillatus var. calcareus (R. W. Butcher) C.D.K. Cook are considered. Brachycentrus subnubilus Curtis has one generation each year, with five larval instars. Larvae favour sites beneath the weedbeds for most of the year, migrating to more exposed areas in the spring to pupate. Pupation, emergence, oviposition and eclosion are completed in just over two months. Eggs are usually laid on submerged marginal vegetation but any semi-submerged object may be utilized. Larvae appear to disperse mainly as first instars; in this they are likely to be seriously affected by weedcutting activities. Initial development is rapid to fourth instar. Larvae overwinter as fifth instars. Sex ratios show a predominance of males.     

Interactions between the Parasitoid Rhopus nigroclavatus (Ashmead) (Hymenoptera: Encyrtidae) and its Mealybug Hosts Tridiscus sporoboli (Cockerell) and Trionymus sp. (Homoptera: Pseudococcidae)

This research investigated age-class-specific parasitism rates of the buffalograss mealybugs Tridiscus sporoboli (Cockerell) and Trionymus sp. by Rhopus nigroclavatus (Ashmead) (Hymenoptera; Encyrtidae), size class preference of this parasitoid, and mealybug–parasitoid interactions through choice and no-choice studies. In the no-choice studies, the mean rates of parasitism by R. nigroclavatus were 45, 20, 0, and 0%, respectively, for mealybugs adult female, third and fourth instars, first and second instars, and eggs. Choice studies indicated that rate of parasitism increased with host size. The mean rates of parasitism on mealybugs in the choice studies were 100% for adult females, 24% for third and fourth instars, 0% for first and second instars, and 0% for eggs. A second set of choice studies investigating mealybug/parasitoid behavior revealed that R. nigroclavatus oviposits in all post-egg mealybug age classes, but first and second instars were less often parasitized than older mealybugs.     

Age-specific Chaoborus predation on rotifer prey

SUMMARY. 1. This is the first study to examine predator-prey interactions between Chaoborm instars and rotifer prey. The predatory behaviour of instars I–III of Chaoborus pimctipennis and the diet selectivity of instars I—IV feeding on rotifers were examined in the laboratory. Prey used in direct observations of predatory behaviour included a variety of rotifers (Symhacta pectlnata, S. ohUmga, Polyarthra remata, Asplanchna girodi, Keratella crassa, spined and unspined forms of Keratella cochlearis) and two crustaceans (Bosmitia longirostris, Mesocyclops edax nauplii. 2. In general, strike efficiencies (percentage of strikes resulting in inges- tion) increased in successive instars I—III. Early instar (I and II) strike efficiencies were low when compared with other invertebrate predators. For a given instar. mean prey handling times varied among prey species more than strike efficiencies. Mean handling times for small, soft-bodied rotifers were lowest and those for wide, hard-bodied prey were highest. 3. Instar I exhibited significantly greater selectivity for the small, soft- bodied S. obUmga than for the larger S. pectinata, hard-bodied K. crassa, and spined and unspined forms of K. cochlearis. Instars II—IV positively selected both the large and small Symhaeta species over all Keratella species. The relationship between Chaobortts selectivity and prey value (weight of prey per unit handling time) can be described by a power function. Ingestion rates of rotifers by older instars (III and IV) are among the highest reported for invertebrate predators. 4. Rotifer vulnerability to Chaoborus predation probably depended on rotifer cuticle texture, body width, and hydrodynamic disturbances. Spined rotifers were not necessarily protected from Chaoborus predation because Chaohorus can manipulate and swallow them. Giguere et al.'s 1982) encounter rate model must be modified to predict encounter rates of slow-moving rotifer prey with Chaohorus.     

The acute toxicity of cadmium to different larval stages of Chironomus riparius (Diptera: Chironomidae) and its ecological significance for pollution regulation

Summary Acute toxicity tests using a static-with-replacement testing procedure were carried out with the four larval instars of the freshwater detritivore Chironomus riparius (Meigen). Median lethal concentrations (10, 24, 48 and 96 h LC50's) indicated great differences in sensitivity to cadmium between instars. Larvae became more tolerant with increasing age, the most resistant stage (fourth instar) having a 24 h LC50 of 2,400 mg Cd l^-1, approximately 950 times greater than the corresponding value of 2.1 mg Cd l^-1 recorded for the most sensitive (first instar) stage. These observations are discussed in relation to the establishment of water quality standards.     

Food and feeding of aquatic larvae of the midge chaoborus flavicans (meigen) (diptera : chaoboridae) in the laboratory

Laboratory observations on the feeding of fourth instar larvae of Chaoborus flavicans (Meigen) show that features relating to the prey, such as swimming behaviour, size and shape, play a major role in determining the kind of food eaten by the predator.In tests with two interacting limnetic cyclopoid copepods, Cyclops abyssorum out-competed Mesocyclops leuckarti when Chaoborus was absent but did not do so when Chaoborus was present.     

Biological control of Tipula paludosa (Diptera: Nematocera) using entomopathogenic nematodes (Steinernema spp.) and Bacillus thuringiensis subsp. israelensis

Tipula paludosa (Diptera: Nematocera) is the major insect pest in grassland in Northwest Europe and has been accidentally introduced to North America. Oviposition occurs during late August and first instars hatch from September until mid-October. Laboratory and field trials were conducted to assess the control potential of entomopathogenic nematodes (EPN) (Steinernema carpocapsae and S. feltiae) and Bacillus thuringiensis subsp. israelensis (Bti) against T. paludosa and to investigate whether synergistic effects can be exploited by simultaneous application of nematodes and Bti. Results indicate that the early instars of the insect are most susceptible to nematodes and Bti. In the field the neonates prevail when temperatures tend to drop below 10 °C. S. carpocapsae, reaching >80% control, is more effective against young stages of T. paludosa than S. feltiae (<50%), but the potential of S. carpocapsae might be limited by temperatures below 12 °C. Mortality of T. paludosa caused by Bti was not affected by temperature even at 4 °C but the lethal time increased with decreasing temperatures. Synergistic effects of Bti and EPN against T. paludosa were observed in 3 out of 10 combinations in laboratory assays but not in a field trial. The potential of S. carpocapsae was demonstrated in field trials against early instars in October reaching an efficacy of >80% with 0.5 million nematodes m⁻² at soil temperatures ranging between 3 and 18 °C. Results with Bti were strongly influenced by the larval stage and concentration. Against early instars in autumn between 74 and 83% control was achieved with 13 kg ha⁻¹ Bti of 5,700 International Toxic Units (ITUs) and 20 kg ha⁻¹ of 3,000 ITUs. Applications in spring against third and fourth instars achieved between 0 and 32% reduction. The results indicate that application of Bti and nematodes will only be successful and economically feasible during the early instars and that the success of S. carpocapsae is dependent on temperatures >12 °C. Synergistic effects between S. carpocapsae and Bti require more detailed investigations in the field to determine maximal effect.     

Leptodora kindti: efficient predator and preferred prey item in Neusiedler See,Austria

In the Neusiedler See, Leptodora kindti appeared in the plankton in April/May; the growing season lasted until November. Its optimum conditions prevailed during the summer months, when densities of >500 ind. m^–3 were recorded. First, males appeared by the mid/end of August, while maximum resting egg production occurred in September. At temperatures below 10 °C the population declined.At high population levels, L. kindti caused suppression of its preferred prey item, the juveniles of Diaphanosoma mongolianum. Twenty-five to 45% of juvenile Diaphanosoma can be eliminated by Leptodora. This predation impact is a pronounced but short-term event. During the periods of high Leptodora densities, a significant relationship between Leptodora predation and the death rate of Diaphanosoma is found.The older developmental instars of L. kindti are themselves endangered by the impact of planktivorous fish (i.e. Pelecus cultratus, Alburnus alburnus), which show a preference for Leptodora larger than 6 mm. Frequently, elimination by fish corresponds with high death rates of Leptodora. During the periods when adult Leptodora occurred in numbers >50 ind. m^–3, 55% of the variance connected with its death rate can be attributed to fish predation. The success of this population of L. kindti is discussed within the framework of predator-prey interactions.     

Stage and age-specific predation in four aphidophagous ladybird beetles

In the present study, stage- and age-specific predatory patterns and nutritional ecology attributes of four co-occurring aphidophagous ladybirds, namely, Coccinella septempunctata (L.), Coccinella transversalis F., Cheilomenes sexmaculata (F.) and Propylea dissecta (Mulsant) on pea aphid, Acyrthosiphon pisum (Harris) were investigated. Despite their lower consumption rates, smaller ladybirds (C. sexmaculata and P. dissecta) and earlier instars were efficient converters of aphid prey with higher relative growth rates, over the respective larger ladybirds (C. septempunctata and C. transversalis) and fourth instars. The consumption rates of pea aphids by the larvae increased with stage, but with a decline in the later part of the fourth instars, just prior to pupation. Adult females of early age groups (between 2 and 7 days) had higher consumption rates than later age groups in all the four species. Further in adult females, consumption rates decrease with increase in age, but without significant changes in conversion efficiencies and relative growth rates.     

Chaoborus crystallinus predation on Daphnia pulex: can induced morphological changes balance effects of body size on vulnerability?

Juvenile Daphnia pulex form neckteeth in reponse to chemicals released by predatory Chaoborus crystallinus larvae. Formation of neckteeth is strongest in the second instar followed by the third instar, whereas only small neckteeth are found in the first and fourth instar of experimental clones. Predation experiments showed that body-size-dependent vulnerability of animals without neckteeth to fourth instar C. crystallinus larvae matched the pattern of neckteeth formation over the four juvenile instars. Predation experiments on D. pulex of the same clone with neckteeth showed that vulnerability to C. crystallinus predation is reduced, and that the induced protection is correlated with the degree of neckteeth formation. The pattern of neckteeth formation in successive instars is probably adaptive, and it can be concluded that neckteeth are formed to different degrees in successive instars as an evolutionary compromise to balance prediation risk and protective costs.