Migrations of haemoglobin-rich Daphnia longispina in a small,steeply stratified,humic lake with an anoxic hypolimnion

Migrations of Daphnia longispina were studied in a small humic lake with an exceptionally shallow oxic epilimnion. Horizontal distributions showed clear avoidance of the shoreline, which might be explained by the lower density of predators (Chaoborus sp. and Notonecta sp.) in the central parts of the lake. In early summer all size classes of D. longispina exhibited upward nocturnal vertical migration, descending to the upper hypolimnion in daytime. Later in summer, when the nocturnally migrating Chaoborus sp. had grown large enough to graze on small Daphnia, the latter seemed to shift towards twilight migration. However, large Daphnia individuals showed no synchronized migration; rather their bimodal vertical distributions suggested asynchronous vertical migration. Large individuals showed a particular tendency to concentrate near to the oxycline, close to the dense phytoplankton and bacteria populations in the upper part of the anoxic hypolimnion. According to vertical trap experiments, large D. longispina visited the anoxic hypolimnion and might harvest its abundant food resources. The high haemoglobin content of large individuals seems a specific adaptation to allow access to low oxygen water and hence to maximize grazing potential, in both epi- and hypolimnion, and minimize predation pressure. By staying predominantly in cooler water near the oxycline, Daphnia might also minimize its energy consumption to adjust to low food availability while sustaining a sufficiently high population density to exploit those unpredictable short periods with abundant food which are common in small headwater lakes. It is suggested that migrations of zooplankton are a complex behavioural adaptation which may not be explained by any single factor. In humic lakes with shallow stratification, vertical migrations seem to offer particularly high potential advantages, because of the short distances between dramatically different environments in the water column. In further studies more emphasis should be placed on migrations of individuals rather than populations, and migrations should be considered as a dynamic part of the structure and function of the whole planktonic ecosystem.     

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.     

Evaluation ofChaoborus predation on natural populations of herbivorous zooplankton in a eutrophic lake

Hydrobiologia - The effect ofChaoborus predation on natural zooplankton communities was tested in enclosure experiments during June, July and August in the eutrophic Frederiksborg Slotssø,...     

Growth of Daphnia longispina L. in a polyhumic lake under various availabilities of algal,bacterial and detrital food

The availability and importance of food sources for growth of Daphnia longispina L. from a highly coloured fishless lake with anoxic hypolimnion were assessed by combining in situ and laboratory experiments. In in situ experiments populations were enclosed in tubes with natural temperature stratification and with or without anoxic hypolimnion. In the laboratory experiments the importance of food source (littoral zone vs pelagic epilimnion) was assessed by enclosing moss thalli and a natural zooplankton population in a large-scale flow-through system supplying food for experimental Daphnia. Growth of juveniles of Daphnia in epilimnetic water was determined in batch culture experiments and the importance of increasing concentrations of bacteria and algae for their growth and development was investigated with a small-scale flow-through system. Access to the anoxic hypolimnion enhanced the growth of Daphnia by 23–24%. Growth rates in the tubes with anoxic hypolimnion were 0.36 and 0.16 d^–1 in July and August respectively. In tubes without anoxia the corresponding values were 0.29 and 0.13. In batch-cultures the highest growth rate determined was 0.16 and the overall rates were lower than in in situ experiments. In batch culture Daphnia was able to grow in darkness for 10 days with a rate of 0.16. In the large-scale flow-through system Daphnia population fed with littoral water reproduced well despite the low concentration of algae and increased its number by a factor of c. 32 in 10 days. However, the animals were small and net production of Daphnia population thus lower under the littoral influence than in the control treatment. Population could survive and grew slowly on pelagial water processed by a natural zooplankton community and with very little algae left. It is thus possible that bacteria serve as a life-support system enabling the population survival over periods of algal shortage. Small-scale flow-through experiments revealed that Daphnia longispina is able to mature and reproduce on a bacterial diet if the food concentration is high enough and Daphnia on bacterial food could achieve growth rates similar to those on an algal diet. The threshold food level for Daphnia longispina was estimated to be c. 18.5 g C 1^–1. Detrital material is of limited value in nutrition of Daphnia even in a lake where more than 75% of carbon is bound in particulate detritus.     

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.     

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.     

The influence of prey behaviour on prey selection of the carnivorous plant Utricularia vulgaris

The mechanisms underlying differential prey selection of two microcrustaceans by the common bladderwort (Utricularia vulgaris) were studied in the laboratory. Functional response experiments with single prey showed that Utricularia had a higher attack rate coefficient and a longer handling time coefficient with the cladoceran Polyphemus pediculus than with the cyclopoid copepod Eucyclops serrulatus. Observation of predation rate, defined as number of prey eaten per unit time, from direct behavioural observation on single prey species, showed a higher predation rate with Polyphemus than on Eucyclops, at low prey densities. The opposite pattern was found at high prey density. When the two prey were presented simultaneously to the predator, Eucyclops was preferred over Polyphemus. Results from the situation with two prey and some of the results from the direct behavioural observations support field data on the diet of Utricularia, which shows that cyclopoid copepods are selected more frequently than Polyphemus.     

The effect of prey composition and abundance on the predation rate and selectivity ofMysis relicta

The predation rate and selection of specific prey byMysis relicta in Lake Tahoe depends on both the total and the relative densities of prey classes. Functional responses of mysids to changes in the density of two species of calanoid copepod prey were curvilinear in laboratory experiments. In the field and in 2-prey treatments, preference was usually positive forEpischura and negative forDiaptomus. However, the preference forEpischura was greatest at the lowest combined densities of prey in the laboratory and decreased at densities >100Epischura / m3 in the field. Several hypotheses to explain the inverse relationship between selectivity and prey density are discussed. Field assemblages of prey vary vertically, horizontally and seasonally. Potential effects of this hetrogeneity on both predator and prey populations are examined.     

Influence of predation by fish and water turbidity on a Daphnia gessneri population in an Amazonian floodplain lake,Brazil

The population behavior of Daphnia gessneri Herbst, 1967 in a floodplain lake (Lago Grande) of the lower Rio Solimões was investigated between April 1979 and March 1980 with regard to 1) predation by the fish called tambaqui (Colossoma macropomum, Characidae), 2) water level fluctuation and 3) water transparency. Zooplankton density samples were collected at two sites near mid-lake, where water depth and Secchi disc transparency were measured. In addition, qualitative samples of zooplankton and fish collections were taken at several sites in the adjacent floodplain areas. The author concludes that fluctuations in Daphnia gessneri populations correlate most with intense predation by fish and water turbidity.     

Studies on functional response and prey selection using zooplankton in the anostracan Chirocephalus diaphanus Prevost, 1803

Freshwater cladocerans and rotifers were used as prey to study functional response and prey selection by adult females of Chirocephalus diaphanus under laboratory conditions. For functional response studies, we offered three rotifer species (Brachionus calyciflorus, B. patulus and Euchlanis dilatata) and three cladoceran species (Alona rectangula, Ceriodaphnia dubia and Moina macrocopa) at various densities ranging from 0.5 to 16 ind. ml^–1. We found increased zooplankton consumption with increasing prey density but beyond 4 ind ml^–1 cladocerans and 8 ind. ml^–1 rotifers, the number of animals eaten plateaued. In general, C. diaphanus consumed fewer large prey (cladocerans) and many more smaller zooplankton (rotifers). For prey selection experiments, we used B. calyciflrous, B. patulus, C. dubia and M. macrocopa, offered at the ratio of two rotifers: one cladoceran and at three prey densities (total zooplankton numbers: 3, 6 and 12 ind. ml^–1). Prey selectivity patterns followed the functional response trends. In general, regardless of prey types, with an increase in the available zooplankton, there was an increase in the number of prey consumed. At any given prey density, C. diaphanus consumed higher numbers of rotifers than cladocerans. Among the prey offered, B. patulus and M. macrocopa were positively selected. Results are discussed in light of possible control of zooplankton by anostracans in temporary ponds.     

Diel horizontal migration and swarm formation in Daphnia in response to Chaoborus

Dense swarms of Daphnia longispina (up to 4000 animals l^–1) were recorded along the littoral zone in a lake where Chaoborus flavicans is considered the main predator. D. longispina coexisted with D. pulex, but there were no D. pulex in the littoral swarms. Swarms were less dense at night (about 1/10 the density), and D. longispina exhibited diel horizontal migrations by aggregating in the littoral during the day and spreading out at night. Laboratory experiments showed that Chaoborus capture efficiency on juvenile daphnids was higher in the light compared to darkness, and that Daphnia exhibited a behavioural response to water that had previously contained Chaoborus. We conclude that predation from Chaoborus can be an important factor affecting the distribution patterns of Daphnia observed in this lake. The behavioural experiments indicated that this influence might be partly mediated by chemical agents.     

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.     

Progress towards characterization of a predator/prey kairomone: Daphnia pulex and Chaoborus americanus

Predatory larvae of the midge Chaoborus americanus induce an antipredator defense (neckteeth) in prey individuals of the cladoceran Daphnia pulex. The signal for presence of predator is a water-soluble chemical. We provide evidence that this kairomone originates in the intestinal tract of the predator. The active compound is an organic molecule of intermediate polarity which is heat stable and partially destroyed by acid and base digestion. It is stable to digestion by the general peptidase Pronase. Hydroxyl groups, but not primary amines, carbonyls or thiols are essential to activity. Low-pressure liquid chromatography on a reverse-phase silica gel (Amicon Matrex C-18) column suggests there may be more than one active component.     

The effects of Chilomonas on the life history traits of [-2pt] Daphnia longispina under semi-natural conditions and the implications for competition in the plankton

Assuming that heterotrophic flagellates (H.F.) can sustain cladoceran life cycles, particularly at periods of low food and high detritus conditions, the growth, reproduction and life span of Daphnia longispinawere studied under conditions of summer food limitation. They were fed both natural resources and natural food enriched with a culture of the colourless Chilomonas (ovoid cell, 8 × 25 m). Four H.F. morphotypes occurred in the natural water and the first experiments with Daphnia, showed that the cladoceran would most easily ingest those of 5 m–10 m, while the addition of Chilomonas severely depressed the H.F. of 2 m. The capability of this flagellate to ingest small H.F. was confirmed using fluorescent particles ranging from 0.94 to 3.95 m. As a consequence, Daphnia could control the abundance of H.F., but also compete with the largest morphotypes. To study the influence of the H.F. on the life span of Daphnia, the cladoceran was acclimated for two generations, before the start of the experiments. In semi-natural conditions, constant temperature, without predators and fed natural water, the summer daphnids achieved smaller sizes, produced fewer offspring and lived for a shorter time than when Chilomonas was added to their diet. Despite the addition of 560 g C l^-1, this was not sufficient to enhance the Daphnia reproduction to the level of the spring population fed natural water. These results clearly illustrate that the heterotrophic flagellates contributed significantly to improving the reproduction and survival of the daphnids. They emphasise the importance of the microbial loop as a link to larger consumers, particularly when unpalatable or nutritionally inadequate algae are dominant, which is often so in lakes during summer.     

Patterns of prey selectivity in the cyclopoid copepod Mesocyclops thermocyclopoides

In the shallow and eutrophic subtropical aquatic ecosystems, which it generally inhabits, the omnivorous copepod Mesocyclops thermocyclopoides encounters a wide variety of animal prey types including ciliates, rotifers, and cladocerans. We studied prey selectivity in laboratory-reared adult females of this species given a choice of (i) prey types belonging to different taxa (ciliates, rotifers, cladocerans, and cyclopoid nauplii), and (ii) different prey species within a taxonomic group differing in body size, morphology or behaviour. We also tested the effect of different proportions of prey species on its selectivity. Prey type proportion had no significant effect on selectivity of the copepod, nor was there any evidence of switching based on the relative abundance of prey. Among the ciliate prey species tested, the largest species, Stylonychia mytilus was positively selected regardless of its relative abundance, while the smallest, S. notophora was selected only when its density was higher. Offered a choice of three species of a brachionid rotifer differing in size, the copepod selected the largest of them, Brachionus calyciflorus, and avoided the smallest B. angularis. The evasive rotifer Hexarthra mira was also avoided. When prey choice included three cladoceran species Daphnia similoides, Moina macrocopa and Ceriodaphnia cornuta, the copepod selected the intermediate-sized M. macrocopa regardless of the abundance of the other two species. Although it fed on Mesocyclops nauplii when there was no choice, M. thermocyclopoides avoided them when alternative food was available. In a multispecies prey choice test, the copepod selected predominantly the rotifer B. calyciflorus and the cladoceran M. macrocopa. We suggest that the prey selectivity patterns shown by M. thermocyclopoides are adaptive in that they lead to ingestion of the most profitable prey.     

Estimating predation risk in zooplankton communities: the importance of vertical overlap

In order to estimate predation risk in nature, two basic components of predation need to be quantified: prey vulnerability, and density risk. Prey vulnerability can be estimated from clearance rates obtained from enclosure experiments with and without predators. Density risk is a function of predator density, and the spatial and temporal overlap of the predator and prey populations. In the current study we examine the importance of the vertical component of overlap in making accurate estimates of predation risk from the invertebrate predator Mesocyclops edax on rotifer versus crustacean prey. The results indicate that assumptions of uniform predator and prey densities cause a significant underestimation of predation risk for many crustacean prey due to the coincident vertical migration of these prey with the predator. The assumption of uniformity is more reasonable for estimating predation risk for most rotifer prey.     

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 influence of prey hardness on Daphnia's selectivity for large prey

Two comparable methods were used to study the feeding of four species of Daphnia on large spherical particles which differed in size and hardness. The first method used gut analysis to estimate the selectivities of daphnids feeding in a broad size range of a single particle type, including polystyrene beads (4–60 µm diameter) in the laboratory and Eudorina colonies (10–90 µm) in the field. In the second method, Daphnia of different sizes fed in a mixture of 6.5 µm Chlamydomonas and one of eight test particles. Smaller daphnids were less effective in feeding on large test particles. Nonlinear regression was therefore used to estimate the Daphnia body size at which the clearance rate on a test particle was reduced to 50% of that for Chlamydomonas. The results of both methods show that prey size and hardness are both very important in determining daphnid feeding selectivity. For a given particle size, soft algae (naked and gelatinous flagellates) are more readily ingested than hard algae (diatoms and dinoflagellates), and hard algae are more readily ingested than polystyrene beads. Daphnia can feed effectively on algae that are 2–5 times larger than the largest ingestible bead.     

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.     

Differential prey utilization by the generalist predator Coleomegilla maculata lengi according to prey size and species

Prey utilization by the generalist predator Coleomegilla maculata lengi Timb. (Coleoptera: Coccinellidae) of three sympatric lepidopterous species was quantified in relation with prey size (age) and prey species. Based on optimal foraging theory, we argued that costs associated with the utilization of small and large prey are higher than those of intermediate prey size. As a result, we expected a higher prey consumption rate on intermediate prey size leading to a convex prey consumption curve. Laboratory experiments showed that, within a given prey instar, Coleomegilla maculata lengi preyed more on Plutella xylostella (L.) compared to Artogeia rapae (L.) and Trichoplusia ni (Hübner). Generally, prey consumption rate by Coleomegilla maculata lengi on the three prey species decreased with increasing immature prey size (age). The predation efficacy of Coleomegilla maculata lengi adults and fourth instar larvae was higher compared to younger coccinellids (L2). Although, Coleomegilla maculata lengi showed a higher level of predation on smaller immature prey, we demonstrated that it is not the optimal size range for this predator. As predicted, prey weight consumption rate by Coleomegilla maculata lengi was higher at intermediate prey size leading to a convex prey utilization curve. The beneficial impact of Coleomegilla maculata lengi predation on the host plant was also estimated by using a Protection Index that considers the differential predation caused by the coccinellids and the relative importance of each pest species in terms of plant injury. Coleomegilla maculata lengi has a more significant beneficial impact when it preys on T. ni immatures.