Different Predation Impacts of Two Cyclopoid Species on a Small-sized Zooplankton Community: An Experimental Analysis with Mesocosms

We used mesocosms to analyze predation impacts on the prey populations and prey community structures by two cyclopoid copepod species, the larger Mesocyclops pehpeiensis and the smaller Thermocyclops taihokuensis, who coexist with small-sized herbivorous zooplankton species in a fish-abundant lake. The overall predation impact on the prey populations was stronger for Mesocyclops than for Thermocyclops. Mesocyclops had a strong and less selective impact on the rotifer community but a selective impact on the crustaceans. In contrast, Thermocyclops had a selective predation impact on rotifers but a weak and less selective impact on the crustacean community. As a result, the former predator reduced the diversity of the crustacean community but not the rotifer community, while the latter had an opposite impact on the diversities of the two communities. It has been suggested that fish induce development of a zooplankton community dominated by the small-sized zooplankton species in fish-abundant lakes. Our results demonstrated that cyclopoid copepods altered species composition and diversity of the small-sized zooplankton community in such lakes. Thus, the results have given an important suggestion on the role of the invertebrate predator cyclopoid copepods, which often coexist with fish, that they determine population dynamics and community structures of small-sized zooplankton in fish-abundant lakes.     

Antennule shape and body size of <Emphasis Type="Italic">Bosmina</Emphasis>: key factors determining its vulnerability to predacious Copepoda

Cyclopoid copepods are common in lakes and ponds, and they have a significant predation impact on the communities of the small zooplankton species. To reduce the predation risk, some cladoceran zooplankters develop protuberant (defensive) morphologies in the presence of the copepods. In the case of the small cladoceran Bosmina, they elongate their appendages (antennule and mucrone) and change the antennule morphotype. However, information about the effectiveness of these defensive devices against copepod predation is still insufficient. In our study, to find the compositive effects of these appendages on the vulnerability of Bosmina, we exposed two bosminid species (B. longirostris and B. fatalis) of different body sizes and with appendages of different lengths and shapes to copepod (Mesocyclops) predation. The experiment revealed that the shape of the antennule is a main factor determining the bosminid’s vulnerability to copepod predation and indicated that the protection of the opened ventral carapace must be a key strategy by which Bosmina avoids copepod predation.     

Top-down control in pelagic systems: a role for invertebrate predation

Limnologists have long recognized the importance of predation in freshwater communities. The majority of study of predator effects has involved vertebrate predators, with emphasis on planktivorous fish. Documented effects of planktivorous fish have been so dramatic that manipulations of their populations are seen by many as potential tools in lake management. However, the success of such manipulations is often less than desired due to the ubiquitous complexity of food webs and the pervasiveness of compensatory responses to food web manipulation. Recently, enormous effort has been applied to the Lake Kinneret pelagic food web in effort to reduced the abundance of the planktivorous Kinneret bleak Acanthobrama terraesanctae and thereby increase the biomass of herbivorous zooplankton in the hopes of increasing water clarity. We compared potential predation pressure on Lake Kinneret herbivorous zooplankton by bleak and the other major zooplankton predators in the lake, the cyclopoid copepods Mesocyclops ogunnus and Thermocyclops dybowskii. We found that, despite having much lower biomass, cyclopoid copepods accounted for a greater portion of the predation mortality on herbivorous zooplankton than bleak. Our results suggest that reductions in predation pressure by bleak will not yield subsequent increases in herbivorous zooplankton biomass. Rather, reductions in bleak predation pressure may allow for increases in cyclopoid copepod abundance and thereby a net increase in predation pressure on herbivorous zooplankton.     

Direct and indirect impacts of predation by fish on the zooplankton community: an experimental analysis using tanks

The impact of Pseudorasbora parva, a common zooplanktivorous fish species in Japan, on a zooplankton community was analyzed in experimental tanks, half of which were stocked with the fish. Different zooplankton species showed different responses to the introduction of the fish. In the presence of the fish, the populations of the large cladoceran Ceriodaphnia and the predatory copepod Mesocyclops were reduced, but the population of the herbivorous copepod Eodiaptomus and the small cladocerans Bosmina fatalis and Bosminopsis deitersi increased relative to the controls. The increase of Mesocyclops seen in the control tanks might have suppressed the populations of the small cladocerans, which are vulnerable to invertebrate predation. The results suggest that the population densities of the large prey items preferred by the fish, Ceriodaphnia and Mesocyclops, were controlled directly by fish predation, but the population densities of the smaller and less preferred zooplankton were controlled indirectly through the food-web cascade.     

Effect of the cyclopoid copepod Mesocyclops thermocyclopoides on the interactions between the predatory rotifer Asplanchna intermedia and its prey Brachionus calyciflorus and B. angularis

In many shallow, eutrophic subtropical ponds, brachionid rotifers are common prey of the predatory copepod Mesocyclops thermocyclopoides. The predatory rotifer Asplanchna intermedia, which is itself a potential prey of the copepod, also feeds preferentially on brachionids. We studied in the laboratory the population dynamics of two mutually competing prey species, Brachionus angularis and B. calyciflorus, in the presence of the two predators A. intermedia and M. thermocyclopoides. The experimental design included separate population dynamics studies with one prey–one predator, two prey–one predator, one prey–two predator, and two prey–two predator systems. These combinations were compared with controls, in which both the prey species (B. angularis and B. calyciflorus) were grown separately and in combination with each other. In the absence of any predator, B. angularis generally eliminated the larger B. calyciflorus. Selective predation by the copepod allowed B. calyciflorus to persist longer in competition with B. angularis. Feeding by M. thermocyclopoides on A. intermedia reduced the predation pressure on B. calyciflorus. However, given enough time, the cyclopoid copepod was able to eliminate both the brachionids as well as the predatory Asplanchna.     

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.     

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.     

The Role of the Predaceous Copepod Parabroteas Sarsi in the Pelagic Food Web of a Large Deep Andean Lake

Parabroteas sarsi is a predaceous calanoid copepod that inhabits both shallow temporary fishless ponds and deep fish lakes of Patagonia and Antarctica. The aim of this study was to analyse the effect of P. sarsi on the plankton structure of a deep Andean lake (>100 m depth) and the zooplankton vertical distribution in order to asses a possible vertical refuge of the predatory copepod against visual fish predation. We tested the extent to which the trophic cascade effect of this predator propagates through the food web. We carried out a vertical sampling in Lake Rivadavia (Patagonia, Argentina) in order to assess zooplankton distribution. P. sarsi showed a vertical distribution towards deeper layers of the water column both at midday and at night, indicating that the copepod had an effective refuge against visual predation. Additionally, we carried out both field and laboratory experiments with the presence of P. sarsi. The predator was observed to affect significantly the survival of the copepod Boeckella michaelseni both in laboratory and field experiments. On the contrary, rotifers and adults of Daphnia cf. commutata were not substantially affected by the predator. B. michaelseni mouthparts revealed an omnivorous diet; therefore a broad phytoplanktonic size spectrum could be affected by this copepod. However, no cascade effect was observed due to the presence of P. sarsi despite the decrease of B. michaelseni abundance.     

Cyanobacterial chemical warfare affects zooplankton community composition

1. Toxic algal blooms widely affect our use of water resources both with respect to drinking water and recreation. However, it is not only humans, but also organisms living in freshwater and marine ecosystems that may be affected by algal toxins. 2. In order to assess if cyanobacterial toxins affect the composition of natural zooplankton communities, we quantified the temporal fluctuations in microcystin concentration and zooplankton community composition in six lakes. 3. Microcystin concentrations generally showed a bimodal pattern with peaks in early summer and in autumn, and total zooplankton biomass was negatively correlated with microcystin concentrations. Separating the zooplankton assemblages into finer taxonomic groups revealed that high microcystin concentrations were negatively correlated with Daphnia and calanoid copepods, but positively correlated with small, relatively inefficient phytoplankton feeders, such as cyclopoid copepods, Bosmina and rotifers. 4. In a complementary, mechanistic laboratory experiment using the natural phytoplankton communities from the six lakes, we showed that changes in in situ levels of microcystin were coupled with reduced adult size and diminished juvenile biomass in Daphnia. 5. We argue that in eutrophic lakes, large unselective herbivores, such as Daphnia, are ‘sandwiched’ between high fish predation and toxic food (cyanobacteria). In combination, these two mechanisms may explain why the zooplankton community in eutrophic lakes generally comprise small forms (e.g. rotifers and Bosmina) and selective raptorial feeders, such as cyclopoid copepods, whereas large, unselective herbivores, such as Daphnia, are rare. Hence, this cyanobacterial chemical warfare against herbivores may add to our knowledge on population and community dynamics among zooplankton in eutrophic systems.     

Experimental analysis of the predation impact of the larvae of pond smelt (<Emphasis Type="Italic">Hypomesus transpacificus nipponensis</Emphasis>) on zooplankton populations established in mesocosms

The predation impact of the larvae of pond smelt Hypomesus transpacificus nipponensis on a zooplankton community was studied using mesocosms. The fish significantly depressed the abundances of copepod nauplii and rotifers, especially Hexarthra mira. The vulnerabilities of these prey might be determined by their swimming behavior and population density, suggesting that larval fish selectively prey on zooplankton that have a high encounter rate with the predator. The larvae did not have a negative effect on the densities of cladocerans, but fish predation altered the cladoceran community structure from the dominance of B. longirostris to that of B. fatalis. This result suggests that larval fish predation is an important factor that shifts the species composition of Bosmina in some lakes, the shift occurring in the season when fish larvae are abundant. Our results have shown that predation by the larval fish would control not only the abundance, but also the community structure of the small-sized zooplankton prey.     

Responses of cyanobacteria to herbivorous zooplankton across predator regimes: who mows the bloom?

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Changes in the abundance and composition of cyclopoid copepods following fish manipulation in eutrophic Lake Væng, Denmark

1. The seasonal cycle of cyclopoid copepods during and following an approximately 50% reduction in planktivorous fish biomass was studied in shallow, eutrophic Lake Vaeng, Denmark, from 1986 to 1990. 2. The dominant cyclopoid copepods changed from Cyclops vicinus and Mesacyclops leuckarti during 1986–1989 to M. leuckarti and Megacyclops viridis in 1990. The abundance of cyclopoid copepods gradually increased from 1986 to 1988–89, decreased in autumn 1989 and markedly decreased in 1990. 3. The increase in the abundance of cyclopoid copepods from 1986 to 1988 is attributed mainly to the reduction in fish predation pressure, there being no concomitant increase in edible phytoplankton. The appearance of M. viridis in 1990, and the general decrease in cyclopoid copepod density in autumn 1989 and in 1990, are attributed to the appearance of submerged macrophytes. 4. Temperature, predation and availability of edible phytoplankton appear to determine whether C. vicinus or M. leuckarti dominates the cyclopoid copepod population of eutrophic Lake Væng.     

Freshwater Copepods and Rotifers: Predators and their Prey

Three main groups of planktonic animals inhabit the limnetic zone of inland waters and compete for common food resources: rotifers, cladocerans and copepods. In addition to competition, their mutual relationships are strongly influenced by the variable, herbivorous and carnivorous feeding modes of the copepods. Most copepod species, at least in their later developmental stages, are efficient predators. They exhibit various hunting and feeding techniques, which enable them to prey on a wide range of planktonic animals from protozoans to small cladocerans. The rotifers are often the most preferred prey. The scope of this paper is limited to predation of freshwater copepods on rotifer prey. Both cyclopoid and calanoid copepods (genera Cyclops, Acanthocyclops, Mesocyclops, Diacyclops, Tropocyclops, Diaptomus, Eudiaptomus, Boeckella, Epischura and others) as predators and several rotifer species (genera Synchaeta, Polyarthra, Filinia, Conochilus, Conochiloides, Brachionus, Keratella, Asplanchna and others) as prey are reported in various studies on the feeding relationships in limnetic communities. Generally, soft-bodied species are more vulnerable to predation than species possessing spines or external structures or loricate species. However, not only morphological but also behavioural characteristics, e.g., movements and escape reactions, and temporal and spatial distribution of rotifer species are important in regulating the impact of copepod predation. The reported predation rates are high enough to produce top-down control and often achieve or even exceed the reproductive rates of the rotifer populations. These findings are discussed and related to the differences between the life history strategies of limnetic rotifer species, with their ability to quickly utilize seasonally changing food resources, and adjust to the more complicated life strategies of copepods.     

Effects of the blue-green alga Microcystis aeruginosa on zooplankton competitive relations

Summary Field distribution patterns and laboratory feeding experiments have suggested that blooms of colonial blue-green algae strongly inhibit relatively large-bodied daphnid cladocerans. We conducted laboratory experiments to test the hypothesis that blooms of the colonial blue-green alga Microcystis aeruginosa would shift competitive dominance away from large-bodied daphnid cladocerans toward smaller-bodied cladocerans, copepods, and rotifers. In laboratory competition experiments, increasing the proportion of M. aeruginosa in the algal food supply resulted in a shift from dominance by the relatively largebodied cladoceran Daphnia ambigua to dominace by the copepod Diaptomus reighardi. The small-bodied cladoceran Bosmina longirostris was always numerically heavily dominant over D. ambigua, but its estimated population biomasses were only slightly higher than those of D. ambigua. Daphnia ambigua consistently outcompeted the rotifer Brachionus calyciflorus. Our results demonstrate that blooms of M. aeruginosa can alter zooplankton competitive relations in laboratory experiments, favoring small-bodied cladocerans and copepods at the expense of large-bodied cladocerans. However, contrary to predictions, blooms of M. aeruginosa did not improve the competitive ability of rotifers.     

Seasonal changes in zooplankton abundance in the lower Rhine during 1987–1991

Zooplankton in the River Rhine was surveyed for five years at the Dutch sampling stations, Lobith (German/Dutch border) and Maassluis (at the point of discharge of the river into the North Sea). The zooplankton abundance showed an apparent seasonal pattern at both stations, characterized by low densities during the winter period, and higher densities during the summer period, with a spring peak. Zooplankton was dominated by rotifers at both stations, although during the winter periods the contribution of copepods was considerable. The rotifers were dominated byBrachionus angularis, B. calyciflorus, Keratella cochlearis andK. quadrata; the copepods by cyclopoid nauplii; the cladocerans by small-sized species mainly belonging toBosmina. At Maassluis the relative contribution of copepods was higher than at Lobith. Furthermore, the zooplankton at Maassluis included the speciesEurytemora affinis, characteristic for estuarine conditions. In spring, the rotifer density and water temperature and rotifer density and chlorophylla concentration were positively correlated. Furthermore, both rotifer density and chlorophylla were inversely correlated with discharge. The possible role of environmental factors (water temperature, chlorophyll content, discharge and biotic factors) controlling the river zooplankton dynamics is indicated.     

Food web structure provides biotic resistance against plankton invasion attempts

It is generally accepted that native communities provide resistance against invaders through biotic interactions. However, much remains uncertain about the types of ecological processes and community attributes that contribute to biotic resistance. We used experimental mesocosms to examine how zooplankton community structure, invertebrate predation, and nutrient supply jointly affected the establishment of the exotic Daphnia lumholtzi. We predicted that establishment would increase with declining biomass and diversity of native zooplankton communities and that an invertebrate predator (IP) would indirectly facilitate the establishment of D. lumholtzi due to its relatively long predator-deterring spines. Furthermore, we hypothesized that elevated nutrient supply would increase algal food availability and facilitate establishment. Only when the biomass and diversity of native zooplankton were significantly reduced, was D.␣lumholtzi able to successfully invade mesocosms. Although invertebrate predation and resource supply modified attributes of native zooplankton communities, they did not influence the establishment of D. lumholtzi. Overall, our␣results are consistent with observed population dynamics in invaded reservoirs where D.␣lumholtzi tends to be present only during the late summer, coinciding with historic mid-summer declines in native zooplankton populations. Lakes and reservoirs may be more susceptible to invasion not only by D. lumholtzi, but also by other planktonic species, in the late summer when native communities exhibit characteristics associated with lower levels of biotic resistance.     

Predatory interactions between a cyclopoid copepod and three sibling rotifer species

SUMMARY 1. Cyclopoid copepod predation on rotifers affects the dynamics and structure of zooplankton communities. We address the differential vulnerability of three sympatric rotifer sibling species belonging to the Brachionus plicatilis species complex. These co-occur with their cyclopoid predator, Diacyclops bicuspidatus odessanus .
2. Using video recording and tracking, we analysed the steps in predation including attack distance, attack angle, and rotifer species swimming in the presence and absence of the predator. Our results show the greater vulnerability of B. rotundiformis (the smallest species) to D. b. odessanus predation, which is associated with a high percentage of attacks after contact. Brachionus plicatilis (the biggest species) is the less vulnerable prey, with low percentage of attacks after contact and captures after attacks. Branchionus ibericus , the intermediate sized species, had also intermediate vulnerability.
3. The differential vulnerability provides insight into the coexistence and seasonal succession of these competing rotifer species. Our results show that the competitive superiority of B. rotundiformis may be balanced by its greater vulnerability to copepod predation.     

Prey preference and utilization by Mesostoma lingua (Turbellaria,Rhabdocoela) at a low arctic site

The rhabdocoel Mesostoma lingua is a common inhabitant of rock bluff ponds in the area of Churchill, Manitoba. Unlike most flatworms it has the ability to prey actively on zooplankton. Among the zooplankton, Mesostoma prefers Daphnia pulex, Simocephalus vetulus, and Diaptomus tyrrelli. Larger calanoid copepods are difficult to capture and take much longer to consume. Growth in Mesostoma is maintained when individuals consume 0.5 Daphnia day^–1 and is maximal at 4 Daphnia day^–1. In comparison, Mesostoma maintained growth only when offered the copepod Diaptomus victoriaensis at the rate of 8–10 individuals-day^–1. Mesostoma may be an important agent in structuring pond zooplankton communities in the arctic, producing a shift in dominance away from cladocerans such as Daphnia toward larger calanoid copepods.     

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.