Where to stay by night and day: Size-specific and seasonal differences in horizontal and vertical distribution of Chaoborus flavicans larvae

1. Data on the distributions of pelagic and benthic Chaoborus flavicans larvae were gathered in 1994 and tested for their agreement with the predator avoidance hypotheses. The development of all Chaoborus life stages, as well as the horizontal and vertical distribution in the four larval instars, was followed from May until October. We expected the largest larvae to dwell deeper by day, thus avoiding predation by visually foraging fish.
2. In agreement with this prediction body size increased with daytime depth, and this was true both between and within instars. The migration amplitude consequently increased with larval instar.
3. There was also evidence for horizontal migration, mainly in the third but also in the fourth instar.
4. Along a horizontal transect with increasing depth, locations with many benthic larvae had fewer pelagic larvae. Oxygen concentration was a good predictor of maximum benthic larval depth for most of the season but failed to predict their distribution in autumn.     

Population interaction: A synchronizer for the persistent rhythmicity of Chaoborus larvae

Fifty chaoborid larvae (Diptera: Chaoboridae) were maintained singly in 200-ml. experimental chambers with lake water and sediments. The larvae, entrained in a reversed light-dark cycle-L D 12: 12 (21–09), were monitored at two-hour intervals to observe whether or not the larvae were in the planktonic state. Results indicated that under constant conditions individual larvae varied in the time of movement out of the sediments and subsequent time spent in the planktonic state. A significant difference was observed when the individual activity patterns (i.e. the number of times an individual was observed to be planktonic during the 12-hour observations period) were summated and compared with the population data previously published by LaRow (1968). However, when larvae whose individual activity patterns were recorded, were pooled in a single experimental chamber the activity pattern then observed was identical to that previously reported. It was concluded, that the persistent, diurnal rhythm observed in Chaoborus is most likely a population rhythm and the interaction of the individuals within the population aids in the synchronization of the rhythm.The observation of larvae maintained singly in experimental chambers also enabled one to determine the actual percentage of the benthic population that leave the sediments and become planktonic each night. During the summer months at least 82.8 percent of the fourth instar larvae leave the sediments and enter the water column.     

The interaction of Chaoborus size and vertical distribution determines predation effects on Daphnia

1. Larvae of Chaoborus, the phantom midge, are important pelagic planktivores in many freshwater lakes and ponds. The effect of Chaoborus on its prey depends on its size, especially mouth gape diameter, and vertical migration pattern, which affects predator–prey spatial overlap. These two features vary considerably in different Chaoborus species and instars. In this study, the interacting effects of both Chaoborus size and vertical distribution on population growth of Daphnia pulex was analysed with a field enclosure experiment and a matrix population model. 2. In the field experiment, Daphnia were grown in four replicated treatments that included a control (no Chaoborus) and three combinations of instar III and IV Chaoborus of two species (C. trivittatus and C. americanus). Parameters of the matrix model were based on differences between Chaoborus species and instars in capture and ingestion of Daphnia of differing sizes (prey vulnerability) and in vertical overlap with Daphnia in each treatment (density risk). 3. In comparison with the control, the two treatments containing the smaller, migratory C. americanus showed a significant effect on Daphnia population growth rate, while the treatment containing only the larger, non‐migratory C. trivittatus did not. The model accurately simulated these effects. 4. A Daphnia predation risk model, which uses prey vulnerability and density risk parameters, illustrated the individual and combined effect of the different Chaoborus types on Daphnia. Daphnia have a high prey vulnerability to the large C. trivittatus, but overall predation risk was low because of very little overlap. On the contrary, the smaller C. americanus affects only a small range of Daphnia instars, each with a low vulnerability, yet those instars that were vulnerable had a very high density risk because of an increased overlap. 5. This analysis of Daphnia predation risk parameters with coexisting Chaoborus species strongly supports an integrated approach using both size and vertical distribution to determine the ultimate predation effect on Daphnia.     

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.     

Implications of Chaoborus pupation and ecdysis in cold water

1. Cold water acted differently to delay and lengthen the pupation period for the larvae of two species of the zooplankton predator Chaoborus (Diptera: Chaoboridae). During Chaoborus pupation, the zooplankton community is released from predation, while the dark-coloured Chaoborus pupae are more susceptible to their own predators.
2. Fourth instar larvae of Chaoborus americanus and C. trivittatus , collected from an oligotrophic lake, were reared individually at 5 °C in the dark. Chaoborus americanus was also reared at 9 and 12 °C under spring photoperiod conditions (L : D, 16 : 8 h). Individuals were observed through pupation to emergence (ecdysis) or death.
3.  Chaoborus americanus pupated at 5, 9 and 12 °C with substantial emergence only at 12 °C. In comparison, C. trivittatus emerged at 5 °C. Light was not a necessary cue for pupation and ecdysis, contrary to previous reports. Cold water delayed the onset and lengthened and increased the variability of the duration of pupation.
4. In Shirley Lake, C. americanus pupated in late June–early July while C. trivittatus pupated first in April and again in June–July.
5.  Chaoborus americanus pupae needed a temperature cue to complete ecdysis. The ecdysis temperature threshold helps to explain the difference in pupation timing, and the geographical distribution, of C. americanus and its relatively inflexible life history contrasted with C. trivittatus . Delayed predator pupation in years with low spring temperature can affect the community dynamics of the prey.     

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.     

An experimental study of the effects of nutrient supply and Chaoborus predation on zooplankton communities of a shallow tropical reservoir (Lake Brobo, Côte d'Ivoire)

1. Based on two mesocosm experiments and 10 in vitro predation experiments, this work aimed to evaluate the impact of nutrient supply and Chaoborus predation on the structure of the zooplankton community in a small reservoir in Côte d'Ivoire. 2. During the first mesocosm experiment (M1), P enrichment had no effect on phytoplankton biomass (chlorophyll a) but significantly increased the biomass of some herbivorous zooplankton species (Filinia sp, Ceriodaphnia affinis). During the second experiment (M2), N and P enrichment greatly increased phytoplankton biomass, rotifers and cladocerans (C. affinis, C. cornuta, Moina micrura and Diaphanosoma excisum). In both experiments, nutrient addition had a negative impact on cyclopoid copepods. 3. Larger zooplankton, such as cladocerans or copepodites and adults of Thermocyclops sp., were significantly reduced in enclosures with Chaoborus in both mesocosm experiments, whereas there was no significant reduction of rotifers and copepod nauplii. This selective predation by Chaoborus shaped the zooplankton community and modified its size structure. In addition, a significant Chaoborus effect on chlorophyll a was shown in both experiments. 4. The preference of Chaoborus for larger prey was confirmed in the predation experiments. Cladocerans D. excisum and M. micrura were the most selected prey. Rotifer abundance was not significantly reduced in any of the 10 experiments performed. 5. In conclusion, both bottom‐up and top‐down factors may exert a structuring control on the zooplankton community. Nutrients favoured more strictly herbivorous taxa and disadvantaged the cyclopoid copepods. Chaoborus predation had a strong direct negative impact on larger crustaceans, favoured small herbivores (rotifer, nauplii) and seemed to cascade down to phytoplankton.     

Effects of prey dispersal on predator–prey interactions in streams

1. We studied the effect of mesh size (6 and 3 mm) on interactions between brown trout ( Salmo trutta ) and benthic invertebrates in enclosures placed in a stream in southern Sweden. We also compared how different prey exchange rates affected interactions between trout and invertebrates.
2. Trout had strong impacts on some benthic taxa, and different mesh sizes produced different patterns. Trout affected the abundance of 10 of the 21 taxa examined, six in enclosures with 3 mm mesh and six in enclosures with 6 mm mesh. The abundance of nine of the prey taxa was lower in the presence of trout, only leptocerids were more numerous in the presence of trout.
3. Our measurements of prey immigration/emigration, together with trout diet data, suggest that direct consumption by trout, rather than avoidance behaviour by prey, explains most decreases in prey abundance. There was avoidance behaviour by only two of the twenty-one prey taxa, with trout inducing emigration of the mayflies Baetis rhodani and Paraleptophlebia sp.
4. Trout indirectly increased periphyton biomass in both 3 and 6 mm enclosures. The effect of trout on periphyton was probably due to strong effects of trout on the grazer, Baetis rhodani , Heptagenia sp. and Paralepthoplebia sp.
5. Our results suggest that mesh size, through its effects on exchange rates of prey, may affect interactions between predators and prey in running waters, but that the effects of dispersal and predation on invertebrates are taxon specific.     

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.     

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.     

Diet of intraguild predators affects antipredator behavior in intraguild prey

In two-predator, one-prey systems with intraguild predationand patchily distributed prey, the intraguild prey may facea choice between prey patches with and without intraguild predators.To minimize falling victim to intraguild predation, intraguildprey are expected to perceive cues specifically associated withthe presence of intraguild predators. We investigate whetherintraguild prey avoided intraguild predators and which cuestriggered this behavior in a system composed of plant-inhabitingarthropods. We found that intraguild prey recognized intraguildpredators from a distance, based on their diet: they avoidedodors of intraguild predators that had consumed shared preybut did not avoid odors of intraguild predators that had fedon other diets, including a diet of conspecifics. When intraguildprey were foraging on a patch, detection of intraguild predatorsled to longer periods of immobility and to fewer captures ofthe shared prey. However, intraguild predators that were eitherstarved or had previously consumed intraguild prey posed a higherrisk to intraguild prey than did intraguild predators that hadconsumed the shared prey. We conclude that the cues used byintraguild prey to avoid intraguild predators are associatedwith the circumstances under which they encounter intraguildpredators in the field and not to different degrees of danger.     

The influence of variability in species trait data on community‐level ecological prediction and inference

Species trait data have been used to predict and infer ecological processes and the responses of biological communities to environmental changes. It has also been suggested that, in lieu of trait, data niche differences can be inferred from phylogenetic distance. It remains unclear how variation in trait data may influence the strength and character of ecological inference. Using species‐level trait data in community ecology assumes intraspecific variation is small in comparison with interspecific variation. Intraspecific variation across species ranges or within populations may lead to variability in trait data derived from different scales (i.e., local or regional) and methods (i.e., mean or maximum values). Variation in trait data across species can affect community‐level relationships. I examined variability in body size, a key trait often measured across taxa. I collected 12 metrics of fish species length (including common and maximum values) for 40 species from literature, online databases, museum collections, and field data. I then tested whether different metrics of fish length could consistently predict observed species range boundary shifts and the impacts of an introduced predator on inland lake fish communities across Ontario, Canada. I also investigated whether phylogenetic signal, an indicator of niche‐conservativism, changed among measures. I found strong correlations between length metrics and limited variation across metrics. Accordingly, length was a consistently significant predictor of the response of fish communities to environmental change. Additionally, I found significant evidence of phylogenetic signal in fish length across metrics. Limited variation in length across metrics (within species), in comparison with variation within metrics (across species), made fish species length a reliable predictor at a community‐level. When considering species‐level trait data from different sources, researchers should examine the potential influence of intraspecific trait variation on data derived by different metrics and at different scales.     

Studies on the biology of Chaoborus flavicans (Meigen) (Diptera: Chaoboridae) in a fish-free eutrophic pond,Japan

Population dynamics of Chaoborus flavicans larvae of various instars was studied from November 1986 to December 1987 in a eutrophic, fish-free pond, Japan. First and 2nd instar larvae were observed from late April to late October, indicating a reproductive period of about half a year. C. flavicans overwintered in the 4th instar larvae. In water column samples, total density of all instars was 680–23680 m^-2, and pupal density 0–2600 m^-2; larvae of the 1st, 2nd, and 3rd instars showed 5–6 density peaks in 1987, suggesting that 5–6 generations occur during a year (peaks of the 4th instar larvae were not clear, probably due to their longer development than those of younger instars). In sediment samples, no 1st and 2nd instar larvae were found, 3rd instar larvae were found occasionally but density of the 4th instar larvae was 280–18600 m^-2, and pupal density varied between 0–502 m^-2. Fouth instar larvae accumulated in sediment in the cold season and in the water column in the warm season; high temperature and low oxygen concentration were the most important factors limiting the distribution of larvae in the sediment in summer in the NIES pond. The dry weight of total C. flavicans larvae was 0.08–4.2 g m^-2 in sediment samples and 24–599 μg l^-1 (0.10–2.40 g m^-2) in water column samples. Comparisons of maximum densities in the NIES pond in different years and in waters of different trophic status show that density is generally higher in eutrophic than in oligotrophic habitats. This revised version was published online in July 2006 with corrections to the Cover Date.     

马尾松毛虫幼虫的捕食天敌及其捕食作用的研究

根据林间调查和室内观察资料,分析了江西省万年县不同松林中马尾松毛虫不同发生代别低龄幼虫期的主要捕食天敌种类及数量。结果表明低龄幼虫期捕食无敌有13科31种,其中以蜘蛛类最多,其次为蚂蚁类。不同林型中捕食天敌的种类和数量在年份间代别间均存在一定差异,数量差异尤其显著。在室内研究了几种主要捕食无敌对马尾松毛虫低龄幼虫的捕食作用及其功能反应,结果表明功能反应为S型,由此建立了它们的功能反应模型。     

Unravelling the nocturnal appearance of bogue Boops boops shoals in the anthropogenically modified shallow littoral

In the present study the role of the nocturnal migration of bogue Boops boops shoals to anthropogenically modified shallow littoral locations was examined, evaluating four alternative hypotheses: (1) feeding, (2) reproduction, (3) attraction of B. boops to artificial light and (4) concealment in the darkness related to predation avoidance. All hypotheses apart from predation avoidance were rejected, as B. boops tended to concentrate in shaded locations of wider illuminated areas, a finding not only important concerning fish behaviour, but also with significant management implications.     

Vulnerability of late larval and early juvenile Atlantic herring, Clupea harengus, to predation by whiting,Merlangius merlangus

Laboratory experiments investigated changes in the vulnerability of herring to predation by whiting during growth from <34 to >80mm in length, Premetamorphic herring (<50mm) failed to react to c . 50% of attacks by the predator, but this was reduced to c . 20% in postmetamorphic fish. Premetamorphic herring failing to react had c. 30% probability of survival due to unforced errors by the predator (unsuccessful attacks that did not elicit a reaction by the prey), but these errors did not occur after herring metamorphosis. The loss of the advantage of lower conspicuousness following metamorphosis was counterbalanced by increased reactivity. Antipredator benefits should increase with schooling behaviour (first evident in experiments at c . 50 mm), so predation mortality should decrease in postmetamorphic herring in the sea. Metamorphosis itself would be associated with high predation risk since conspicuousness is enhanced but reactivity and schooling behaviour are still not fully developed.     

Chaoborus predation and delayed reproduction in Daphnia: a demographic modeling approach

I develop a demographic model that examines the impact of Chaoborus predation on the population dynamics and life history of Daphnia. Predation effects are determined through analysis of the various components of the predator-prey interaction (encounter, attack, strike efficiency), and are integrated into a stage-classified matrix population model. The model is parameterized with data from interactions between D. pulex and fourth-instar C. americanus. I test this model with two laboratory experiments that examine population growth of D. pulex under the influence of five different levels of Chaoborus predation. With the exception of a single predation treatment in each experiment, the model accurately predicted the observed reduction in Daphnia numbers with increasing Chaoborus predation. I then use this model to investigate the evolution of delayed reproduction in D. pulex that are exposed to Chaoborus. I ask whether delayed reproduction may evolve in Daphnia that are subjected to Chaoborus predation as a trade-off for the benefits of larger body size. The model predicts that the effectiveness of such a life history trade-off depends on the relative sizes of predator and prey. In some interactions between Chaoborus and Daphnia, increasing Daphnia body length by as little as 5% from base growth trajectories sufficiently increases fitness (by reducing vulnerability to Chaoborus predation) to compensate for the cost of delayed reproduction. In other interactions, however, increased body length provides no benefit to Daphnia (and may even reduce fitness), and selection would act against the evolution of delayed reproduction. This revised version was published online in July 2006 with corrections to the Cover Date.     

Population decline and life‐cycle changes in a phantom midge (Chaoborus flavicans) after introduction of planktivorous fish

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The influence of hunger and predation risk on group cohesion in a pelagic fish, walleye pollock Theragra chalcogramma

Variation in the intensity of schooling behavior in fishes suggests that the benefits of aggregation are balanced by certain costs. We examined the proximity of group members to each other in juvenile walleye pollock, Theragra chalcogramma, under different environmental conditions. Food availability, simulated by a gradient of six ration treatments, had a major influence on group cohesion, with increasing dispersion as food level decreased. Group cohesion also decreased at night relative to daytime levels. Small juveniles (x=53 mm TL) maintained on high rations were highly responsive to the potential threat of a predator, with groups becoming more cohesive and remaining so for up to an hour after the initial threat. A chronic threat (continual presence of predators) resulted in tighter group cohesion than an acute threat (single simulated attack). Small juveniles maintained on low rations were less responsive to predation threats and recovered quickly, supporting the hypothesis that hunger induces risk-taking behavior. Large juveniles (x=149 mm TL) did not change their degree of aggregation in response to either type of predation threat. An overall plasticity in the degree of cohesiveness among group members indicates that walleye pollock are capable of gradually modifying their schooling behavior according to the environmental context.     

The ecological consequences of environmentally induced phenotypic changes

Population dynamics and species persistence are often mediated by species traits. Yet many important traits, like body size, can be set by resource availability and predation risk. Environmentally induced changes in resource levels or predation risk may thus have downstream ecological consequences. Here, we assess whether quantity and type of resources affect the phenotype, the population dynamics, and the susceptibility to predation of a mixotrophic protist through experiments and a model. We show that cell shape, but not size, changes with resource levels and type, and is linked to carrying capacity, thus affecting population dynamics. Also, these changes lead to differential susceptibility to predation, with direct consequences for predator‐prey dynamics. We describe important links between environmental changes, traits, population dynamics and ecological interactions, that underscore the need to further understand how trait‐mediated interactions may respond to environmental shifts in resource levels in an increasingly changing world.