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.     

Do Chaoborus larvae migrate in temporary pools?

A diurnal vertical migration of larvae of Chaoborus punctipennis (Say) was observed in shallow temporary woodland pools in East Texas. In the laboratory, in 153 cm tall columns, the larvae underwent a much greater migration than possible in the shallow pools. We hypothesize that the migratory behavior and transparent body of Chaoborus larvae may have originally evolved in such shallow water habitats. These traits enabled Chaoborus to successfully invade the quite different adaptive zone of predation in the plankton of deep lakes.     

Density-dependent predation ofChaoborus flavicans onDaphnia longispina in a small lake: the effect of prey size

Functional response curves of fourth instar larvae ofChaoborus flavicans preying on two size classes ofDaphnia longispina were examined throughout three summer seasons in a small forest lake. Data for each size class were fitted to Holling's disc equation. The parametersa (attack rate) andTh (handling time) were calculated for each prey size from these curves. Attack rate was greater and handling time was shorter for small (0.77 mm) than for large (1.82 mm)Daphnia. In 1:1 mixture of these prey size classes the predation rates ofChaoborus on smallDaphnia at prey densities above 20 l^–1 were greater than predicted from the single size-class experiments. The observed predation rates on largeDaphnia were lower than predicted at all prey densities. Since both single size-class and two size-class experiments were run during the same period of time the difference in observed and predicted predation rates could not be attributed to seasonal changes in prey preference ofChaoborus larvae. In experiments with a concentrated mixture of lake zooplankton (dominated byD. longispina)Chaoborus preference forDaphnia decreased as prey body size increased. There was no obvious correlation between selectivity coefficients and size-frequency distributions ofDaphnia. When medium-sizedDaphnia were omitted from calculations the preference of small over large prey did not differ significantly from the predictions of the single size-class model.     

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 selection by Chaoborus punctipennis under laboratory conditions

Equal numbers of Diaphanosoma leuchtenbergianum, Daphnia parvula, and Diaptomus pallidus were offered to individual, fourth-instar larvae of Chaoborus punctipennis. When the prey species were presented separately to the larvae, 100% of the Diaphanosoma, 67% of the Daphnia, and 57% of the Diaptomus were consumed. However, when the three species were offered to the predator concurrently, 90% of the Diaphanosoma, 10% of the Daphnia and < 1% of the Diaptomus were eaten. This strong selection for Diaphanosoma was consistent at three prey densities.     

A metalimnetic oxygen minimum indirectly contributing to the low biomass of cladocerans in Lake Hiidenvesi – a diurnal study on the refuge effect

The diurnal vertical migrations of smelt (Osmerus eperlanus), larvae of phantom midge (Chaoborus flavicans) and cladoceran zooplankton in eutrophic Lake Hiidenvesi were studied in order to clarify the factors behind the low zooplankton biomass. In the study area, an oxygen minimum occurred in the metalimnion in the 10–15 m depth. No diurnal fluctuations in the position of the minimum were observed. Cladocerans inhabited the epilimnion throughout the study period and their vertical movements were restricted to above the thermocline and above the oxygen minimum. C. flavicansconducted a diurnal migration. During the day, the majority of the population inhabited the 12 – 15 m depth just in the oxygen minimum, while during darkness they were found in the uppermost 8 m. Smelts started ascending towards the water surface before sunset and reached the uppermost 3 m around 23:00. During daytime, the majority of smelts inhabited the depth of 7–9 m, where the water temperature was unfavourably high for them (18 °C). Smelts thus probably avoided the steep oxygen gradient in the metalimnion, whereas Chaoborusused the oxygen minimum as a refuge against predation. Those smelts that were found in the same water layers as Chaoborusused the larvae as their main prey. The metalimnetic oxygen minimum thus seemed to favour the coexistence of vertebrate and invertebrate predators, leading to a depression of cladoceran zooplankton.     

Mesocosm experiment on the impact of invertebrate predation on zooplankton of a tropical lake

Invertebrate predation on zooplankton was investigated in mesocosms in the shallow tropical Lake Monte Alegre, São Paulo State, Brazil, in the summer of 1999. Two treatments were applied: one with natural densities of prey and the predators Chaoborus brasiliensis and the water mite Krendowskia sp. (Pr+), and another without predators (Pr-). Three enclosures (volume: 6.6 m³ of water per enclosure) per treatment were installed in the sediment of the deepest area of the lake (5.0 m). At the beginning, Chaoborus larvae were present in Pr- enclosures, because of technical difficulties in preventing their entrance, but they virtually disappeared in the course of the experiment. Water mites were almost absent in Pr- enclosures. Chaoborus predation negatively influenced the Daphnia gessneri population, but not the populations of the copepods Tropocyclops prasinus and Thermocyclops decipiens and the rotifers Keratella spp. Death rates of Daphnia were generally significantly higher in the Pr+ treatment; Daphnia densities increased after the disappearance of Chaoborus in Pr-. Copepod losses to predation in the experiment may be compensated by higher fecundity, shorter egg development time, and lower pressure on egg-bearing females, resulting in a lower susceptibility to Chaoborus predation. The predation impact of water mite on microcrustaceans and rotifers in the experiment was negligible.     

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.     

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.     

Ingested prey increase risks of visual predation in transparent Chaoborus larvae

Summary Transparency reduces the chances of detection of large planktonic animals by visual predators. An important constraint on the transparency of planktonic animals may be ingested food which could be seen through the body, thereby increasing the vulnerability of transparent zooplankton to visual predators. To test this hypothesis, we presented fed and un-fed Chaoborus larvae to juvenile coho salmon (Oncorhynchus kisutch). Overall, the presence of prey in the gut of Chaoborus increased their probability of capture by 68%. Predation risks due to the visibility of ingested food increased in proportion to meal size: larvae with nearly full gut were captured about three times faster on the average than larvae which had little food in their gut. Although Chaoborus larvae may be able to reduce this increased predation risk by migrating downward to low light levels, this behavior would reduce feeding opportunities by removing the larvae from surface waters where prey density is generally high. In this way, visual predators may limit the growth and the maximum size that can be achieved by transparent animals.     

Influence of benthic and interstitial processes on nutrient changes along a regulated reach of a large river (Rhône River,France)

An enclosure study was conducted in Ranger Lake in south-central Ontario, Canada from 4 July to 5 August 1997 to determine predation effects of the larvae of the phantom midge fly Chaoboruson the zooplankton community. Zooplankton assemblages were established in 12 enclosures (2 m in diameter, 7.5 m deep). Three densities of fourth-instar Chaoborus trivittatus (0 l^–1, 0.1 l^–1 and 0.5 l^–1) were introduced as predator treatments to the enclosures. Temperature, dissolved oxygen and zooplankton community composition were monitored for six weeks. To determine if the zooplankton community composition changed, a repeated measures multivariate analysis was performed on percent biomass of Bosmina and calanoid copepods. There were no significant differences in mean taxon percent biomass among predator treatments. There were significant differences in mean taxon percent biomass between water layers (epilimnion and metalimnion). There were also significant differences in lengths of Bosmina and calanoid copepods among predator treatments at the end of the experiment. Crop content analysis of C. trivittatusshowed that Bosmina constituted 88–98% of the prey items found in the crops. These results demonstrate that the use of deep enclosures, a Chaoborus species which vertically migrates, and lower natural densities of Chaoborus may provide prey with an important natural refuge from predation and so allow a more accurate determination of the predation impact of Chaoborus trivittatusin temperate lakes where fish control Chaoborus densities.     

Predation and refugia: implications for Chaoborus abundance and species composition

1. Previous studies have suggested that the occurrence of larval Chaoborus in lakes may be affected by fish predation, pH, elevation, temperature, nutrient level, water transparency and interspecific competition, but so far, a detailed statistical evaluation of these findings has not been performed. 2. The aim of this study was to apply regression and ordination techniques to a large data set of 56 lakes in order to test which variables related to lake morphology, water chemistry, and fish predation determine (1) the abundance of individual Chaoborus species and (2) their species composition. 3. Individual Chaoborus species were influenced by very different sets of environmental factors. Nutrient levels positively affected the largest species, Chaoborus americanus, which was restricted to fishless lakes. Abundance of the smallest and most transparent species, C. punctipennis, seemed to be controlled more by the larger Chaoborus species than by fish. Larger chaoborids required low water clarity in order to co‐exist with fish, probably to increase refuge availability. Generally, small lakes (for C. flavicans/C. trivittatus) and shallow lakes (for C. punctipennis) supported higher abundances of Chaoborus.     

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     

Habitat segregation and interactive effects of multiple predators on a prey assemblage

1. In a field experiment we examined the interactive effects of two common predators of zooplankton, bluegill sunfish (Lepomis macrochirus) and Chaoborus spp. on the growth rate and habitat use of three congeneric prey species (Daphnia). Bluegill and Chaoborus both consume Daphnia, but bluegill also prey on Chaoborus. The prey species, Daphnia pulicaria, D. rosea and D. retrocurva, differed in body size and vertical distribution. We expected the largest species, D. pulicaria, to be most vulnerable to fish predation and the smallest species, D. retrocurva, to be most vulnerable to Chaoborus predation. 2. As we expected, the population growth rate of D. pulicaria was significantly reduced by fish. However, Chaoborus also significantly reduced the growth rate of this species. No significant interaction effect was detected, indicating that the effect of these predators was additive. The growth rates of D. rosea and D. retrocurva were significantly reduced by Chaoborus, but a significant interaction effect indicated that the effect of Chaoborus was stronger in the absence of fish than when fish were present. Therefore the impact of Chaoborus and fish on D. rosea and D. retrocurva was non-additive. The interactive effect of the two predators on D. retrocurva was greater in magnitude than on D. rosea. 3. In the absence of predators, the three Daphnia species showed no differences in mean habitat depth between day and night. Both predators significantly affected diel habitat use of D. pulicaria and D. rosea. Fish caused both of these Daphnia species to move deeper during the day, whereas Chaoborus caused Daphnia to move into shallower water at night. Daphnia retrocurva tended to migrate upwards at night in all predator treatments, but no significant differences in migration were observed among the predator treatments. The effects of predators on habitat use were not interactive for any prey species. 4. Our results suggest that body size, habitat use and the diel migratory response to predators are important factors mediating the interactive effects of multiple predator types on zooplankton.     

The effect of prey type and density on the foraging efficiency of juvenile perch, (Perca fluviatilis)

The foraging efficiency of juvenile perch (Perca fluviatilis), feeding on two types of prey, was studied in laboratory experiments. Waterfleas (Daphnia magna) and phantom midge larvae (Chaoborus flavicans) were offered in a range of densities, either separately or combined. Perch fed more efficiently on each prey type separately than when both were mixed. Foraging efficiency decreased with an increase of mixed prey density with both prey types present in equal numbers, but also when the proportion of Chaoborus increased. This could be caused by the existence of different hunting techniques, each of which is fully efficient in the presence of one prey type only. In the presence of two prey types, the predator constantly has to switch from one hunting technique to another.     

Biotic resistance of impact: a native predator (<Emphasis Type="Italic">Chaoborus</Emphasis>) influences the impact of an invasive predator (<Emphasis Type="Italic">Bythotrephes</Emphasis>) in temperate lakes

Introduced predators have caused some of the largest documented impacts of non-native species. Interactions among predators can have complex effects, leading to both synergistic and antagonistic outcomes. Complex interactions with native predators could play an important role in mediating the impact of non-native predators. We explore the role of the native predator context on the effect of the introduced predatory cladoceran Bythotrephes longimanus. While post-invasion impacts have been well described, studies have largely ignored the role of native predators. We used a field mesocosm experiment to determine whether Bythotrephes’ impact on prey communities is influenced by the presence of the ubiquitous native predatory insect larvae Chaoborus. The two predators exhibited niche complementarity as no change in total zooplankton prey abundance was detected across predator treatments. Rather, copepod abundances increased with decreasing abundances of Chaoborus, while cladocerans decreased with increasing abundances of Bythotrephes. Thus, the replacement of Chaoborus with Bythotrephes led to changes in the overall community structure of the zooplankton prey, but had little effect on prey total abundance. More interestingly, we found evidence of biotic resistance of impact, that is, the impact of Bythotrephes on the cladoceran community was altered when the two predators co-occurred. Specifically, the predation effect of Bythotrephes was more restricted to the shallower regions of the water column in the presence of Chaoborus, leading to a reduced impact on deeper dwelling prey taxa. Overall, our results demonstrate that the native predator context is important when trying to understand the effect of non-native predators and that variation in native predator abundances and assemblages could explain variation in impact across invaded habitats.     

Zooplankton community structure driven by vertebrate and invertebrate predators

Summary A zooplankton community was established in outdoor experimental ponds, into which a vertebrate predator (topmouth gudgeon: Pseudorasbora parva) and/or an invertebrate predator (phantom midge larva: Chaoborus flavicans) were introduced and their predation effects on the zooplankton community structure were evaluated. In the ponds which had Chaoborus but not fish, small- and medium-sized cladocerans and calanoid copepods were eliminated while rotifers became abundant. A large-sized cladoceran Daphnia longispina, whose juveniles had high helmets and long tailspines as anti-predator devices, escaped from Chaoborus predation and increased. In the ponds which had fish but not Chaoborus, the large-sized Daphnia was selectively predated by the fish while small-and medium-sized cladocerans and calanoid copepods predominated. In the ponds containing both Chaoborus and fish, the fish reduced the late instar larvae (III and IV) of Chaoborus but increased the early instar larvae (I and II). Small- and large-sized cladocerans were scarcely found. The former might have been eliminated by predation of the early instar larvae of Chaoborus, while the latter was probably predated by fish. Consequently, the medium-sized cladocerans, which may have succeeded in escaping from both types of predator, appeared abundantly. The results suggest that various combinations of vertebrate and invertebrate predators are able to drive various kinds of zooplankton community structure.     

Impact of Chaoborus predation upon the structure and dynamics of a crustacean zooplankton community

Summary During ice-free seasons of 1975, 1977 and 1978, replicated experimental alteration of spring densities of predatory Chaoborus larvae inside 20–27 m³ enclosures in a fishless oligotrophic lake had relatively small, but significant, short-term effects upon prey species abundances. Enhancement of predator densities generally had greater numerical effects relative to controls than did complete removal of predators. With the exceptions Diaphanosoma and Bosmina under artificially elevated Chaoborus densities, numerical effects on prey species did not persist for more than a few weeks after midsummer in these 3 years. During cooler 1976, however, much larger Chaoborus effects in May and June persisted into September. Low temperatures and small initial population densities slowed population growth and tended to increase the proportion of each species' recruitment lost to Chaoborus predation. As water temperatures increased during 3 of 4 summers, rapid juvenile development and compensating increase in adult fertility generally permitted most prey species to escape regulation by these large, univoltine and semivoltine predators. With growing prey population size, declining food levels suppressed crustacean fecundity in July and August, thereby permitting predatory losses to climb again to substantial fractions of the reduced prey recruitment. Nevertheless, prey densities in predator-free and control or predator-enhanced enclosures differed little from July through September (except in 1976). Thus, summer population growth of most prey species seemed more limited by food shortages than by predators per se. Growth of individual zooplankters was affected by food availability during critical periods in July and August in all years, and Chaoborus predation seemed to influence the timing of this food limitation in at least 2 of the 4 years     

Consumption and utilization of a soyflour-wheat germ diet by larvae of the tobacco budworm parasitized by the tachinidEucelatoria sp

The number ( \(\bar X\) =2.4) ofEucelatoria sp. maggots that completed development in 4th- or 5th-instar larvae of the tobacco budworm (TBW),Heliothis virescens (F.), was significantly greater (P<0.05) than the number ( \(\bar X\) =1.2) that completed development in 3rd-instar larvae. Maggot development time decreased with increasing number of maggots per host larva. It also decreased with advancing larval instars. The range was 6.9±1.1 days in early 3rd-instar TBW larvae and 5.0±0.8 days in early 5th-instar TBW larvae. Unparasitized 3rd- or 4th-instar TBW larvae consumed significantly more food than did similar aged larvae parasitized byEucelatoria sp., but larvae parasitized during the early 5th-instar consumed more food than did similar aged unparasitized larvae. Consumption by 4th- or 5th-instar larvae increased significantly as maggot densities increased from 1 to 3 per host larva, but decreased at a density of 4 or more maggots per host larva. Although body weight gain and consumption were both significantly reduced 48 and 120 h after parasitization of late 3rd-instar larvae (6 days old), the approximate digestibility (AD) value was significantly greater for parasitized than for unparasitized larvae. Unparasitized larvae were more efficient in converting digested food to body substance (ECD) than parasitized larvae, but the efficiency in conversion of ingested food to body substance (ECI) was similar for both parasitized and unparasitized larvae.     

The influence of zooplankton defenses on prey selection by young Chaoborus larvae

Prey exhibiting a variety of morphological and behavioral antipredatordefenses were examined for susceptibility to first through thirdinstar Chaoborus americanus larvae. Synchaeta sp. was highlyvulnerable to these predators due to high encounter rates, itsrelatively soft cuticle, and lack of a precontact defense. Keratellatestudo exhibited postcontact defenses and was more susceptibleto predators than were prey of comparable size that exhibitedprecontact escape behaviors (Hexarthra mira and Diaptomus birgeinauplii). Ontogenetic changes were observed in the escape responseof D.birgei and the foraging success of C.americanus larvae.Chaoborus was observed to prey on the protists Bursaria andVolvox. Precontact defenses were more effective than postcontactdefenses in the prey species examined; however, the relativeeffectiveness of prey defenses cannot be generalized by preygenus nor by the defensive tactics employed. Present address: Department of Entomology, University of California,Riverside, CA 92521, USA