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.     

Implication of the feeding limb morphology for herbivorous feeding in some freshwater cyclopoid copepods

1. The mouthparts of five species of adult planktonic cyclopoid copepods (Cyclops vicinus, C. abyssorum, Acanthocyclops robustus, Mesocyclops leuckarti and Thermocyclopscrassus), in particular the distance between setae and setules of the maxilliped (which can indicate the ability to retain small particles), were compared using electron and light microscopy. 2. The mesh‐sizes of the food‐collection grid formed by these setae and setules ranged between 4.6 and 13.2 μm; the area covered by the grid ranged between 6000 and 32 000 μm². 3. Mesh‐size was not simply correlated with body size. Cyclops abyssorum and M. leuckarti have the coarsest meshes and T. crassus the finest, while C. vicinus and A. robustus were intermediate. 4. The results suggest that cyclopoid copepods are able to retain particles in the size range of nanoplankton and that differences in mesh‐sizes between species may explain differences in the ability to subsist and reproduce on a diet of small algae.     

The influence of food resources on the development, survival and reproduction of the two cyclopoid copepods: Cyclops vicinus and Mesocyclops leuckarti

Laboratory experiments were conducted to determine whether thetwo cyclopoid copepods. Cyclops vicious and Mesocyclops leuckaru.exploit the same food resources. The food requirements of juvenilesof the two cyclopoid copepods were investigated. Moreover, theimportance of algae for the predaceous adults was studied. Naupliiof both M leuckaru and C.vicinus successfully developed intocopepodites when fed the motile algae Chlamydomonas reinhardtii.Chlamydomonas sphaeroides and Cryptomonas sp. Threshold foodconcentrations for naupliar development varied between offeredalgae and between the two cyclopoid species. The food thresholdfor successful naupliar development, when reared on C.reinhardui,was lower for M.leuckarti (0.3 mg C 1^–1) than for C.vicinus(0.5 mg C l^–1) whereas a similar food threshold was foundusing Cryptornonas sp (0.3 mg C ^–1) and C.sphaeroides(<0.2 mg C 1^–1), Naupliar development time was inverselyrelated to food concentration. Food required for copepoditedevelopment differed for the two cyclopoid species. Cyclopsvicinus was able to develop to the adult stage on a pure dietof any one of the three algal species. whereas M.leuckarti requireda prey supply of the rotifer Brachionus rubens. Food composition.i.e. algal species, algal concentration and rotifer abundance,influenced copepodite survivorship of both cyclopoids and wasalways higher in the presence of B.rubens. Under similar foodconditions, mortality was higher for M.leuckarti than for Cvicinus. Mesocyclops leuckaru females were very dependent onanimal food. The predation rate of M.leuckaru was not lowerin the presence of algae. Egg production of M.leuckarti waslow on a pure algal diet and significantly higher when B rubenswas present. The results were used to discuss the life cyclestrategy and the possibility of exploitative competition ofthe two cyclopoid copepods.     

An example of niche partitioning in three coexisting freshwater cyclopoid copepods

Vertical distributions at noon and midnight of copepodites,males and females of three coexisting species of freshwatercyclopoid copepods, Cyclops vicinus (a cold-water form), Thermocyclopscrassus and Mesocyclops leuckarti (warm-water forms), were studiedin Dreiangel Lake, a small eutrophic gravel pit. The largestspecies, C. vicinus, inhabits deep water layers, both at noonand midnight, while the smaller species, T.crassus and M.leuckarti,inhabit upper water layers. In all three species, copepoditesand males live closer to the water surface than females. Insummer, abundance of the smallest species (T.crassus) is highest,although finite birth rates are lowest. The distribution patternsand population characteristics suggest that the vertical distributionof cyclopoid copepods in Oreiangel Lake is mainly governed byfish predation; different temperature tolerances and preferencesof the three species may also be of significance.     

The effect of temperature on the development times of eggs,naupliar and copepodite stages of five species of cyclopoid copepods

The duration times of eggs, combined naupliar instars and of the different copepodite stages of five species of cyclopoid copepods — Acanthocyclops robustus, Cyclops vicinus, Diacyclops bicuspidatus, Mesocyclops leuckarti, and Thermocyclops crassus — were investigated at five different temperatures. The five species can be divided in two groups: two species, C. vicinus and D. bicuspidatus, adapted to cold water conditions and three species, A. robustus, M. leuckarti and T. crassus adapted to warm water conditions. The cold water species showed a faster egg development than M. leuckarti and T. crassus at 5–15 °C. The eggs and instars of the warm water species M. leuckarti tend to develop faster than those of the former two species at higher temperatures. A. robustus showed the shortest egg and instar development at 10–25 °C. The warm water species T. crassus produced no eggs at 10 °C and temperatures below. At higher temperatures (20, 25 °C) the egg and instar duration times were similar or longer than those of the other species. When cultured in total darkness a great part of the C_IV respectively C_V copepodites of the summer forms entered arrest and the percentage of copepodites that showed an arrest of development was highest at lowest temperatures. The present results are compared with data from literature and differences are discussed.     

Spatial distribution of resting stages,rate of emergence from diapause and times to adulthood and to the appearance of the first clutch in 3 species of cyclopoid copepods

The spatial distribution of dormant copepodids of 3 species of cyclopoid copepods — Cyclops vicinus, Mesocyclops leuckarti and Thermocyclops crassus — was studied in 4 small lakes in South Germany. The rate of emergence from diapause and times from the resting stage to adulthood and from adulthood to the appearance of the first clutch was studied at 4 constant temperatures (5, 10, 15, 20 °C) in the laboratory. Resting stages of C. vicinus were always concentrated in the deepest parts of the lakes and were found relatively deep in the mud. M. leuckarti- and T. crassus-copepodids preferred shallow areas in deep lakes but were concentrated in the deep areas in shallow lakes. Copepodids of both species were always concentrated in the uppermost mud layers.Rate of emergence from diapause was strongly temperature-dependent. At high temperatures (20 °C) copepodids of all species under study emerged within 2 weeks. At lower temperatures C. vicinus copepodids showed the highest rate of emergence. At 5 to 10 °C only few M. leuckarti- and T. crassus-copepodids had emerged after the investigation period (7 weeks). Both C. vicinus and T. crassus showed the highest rate of emergence at the natural end of diapause but even at that time only few T. crassus-copepodids emerged at 5 °C. Times to adulthood at 5 °C were shortest in C. vicinus. At higher temperatures this species was passed by M. leuckarti. Times from adulthood to the appearance of the first clutch at 5–15 °C were shortest in C. vicinus. T. crassus produced no clutch at 5 and 10 °C.     

Is mating of copepods associated with increased risk of predation?

In laboratory experiments, we tested whether the cyclopoid copepodCyclops vicinus is more vulnerable to predation by juvenile(young-of-the-year) roach (Rutilus rutilus) and/or larvae ofthe phantom midge Chaoborus flavicans during mating than individualadult copepods of the same species. We also tested whether C.vicinusavoided mating, displayed shorter mating times and/or loweractivity when a predator (Chaoborus) was present, or when exposedto water in which predators (Chaoborus) were previously held.Fish ate the same quantity of pairs in copulation and individualegg-bearing females but significantly fewer females withouteggs. Chaoborus ate similar quantities of pairs and femalesbut fewer males. Pairs in copulation and egg-bearing femaleswere detected earlier by fish than individual non-ovigerousfemales or males, and pairs in copulation were captured mosteasily by fish. Cyclops vicinus mated less often when Chaoboruslarvae were present than in water in which Chaoborus was previouslyheld, or in pure tap water. However, predator presence or exposureto predator-treated water had no effect on (i.e. did not reduce)mating duration. Activity of C.vicinus pairs during copulationwas significantly lower when a predator was present, or in predator-treatedwater, than in pure tap water, and activity of pairs decreasedwith increasing activity of the predator. Although our experimentswere carried out under artificial conditions which cannot betransferred to the complex conditions in the field, they suggestthat mating in copepods is dangerous and that copepods haveevolved particular strategies to reduce the risk of predationduring mating.     

The seasonal cycle of Thermocyclops crassus (Fischer, 1853) (Copepoda: Cyclopoida) in a shallow,eutrophic lake

The seasonal cycle of Thermocyclops crassus was studied from 1985 to 1987, in the Gronne, a shallow, productive lake. T. crassus was present from late April to early October, while water temperature was above 10 °C. It produced three generations per year. Population peak was usually reached by the second generation, in August. Abundance was positively correlated with water temperature. Females carried 18.3 to 32.3 eggs on average. Reproduction rates were highest in July and August. Sex ratio was low, as females generally outnumbered males. Between October and April copepodites 4 went in diapause, predominantly in the deepest part of the lake and 0–4 cm deep in the bottom mud. Variations in body size were low across the year. T. crassus coexisted with the cyclopids, Cyclops vicinus and Mesocyclops leuckarti. In 1985 and 1986, abundances of M. leuckarti were low, while those of T. crassus and C. vicinus were high. In 1987 lower water temperatures, caused by cold weather, resulted in a marked decrease of the population density of T. crassus, while the abundance of M. leuckarti increased, and density of C. vicinus remained high. Comparison of the egg duration times of T. crassus, C. vicinus and M. leuckarti showed that at 15 °C and below T. crassus may be outcompeted by its comparatively longer egg development times. However, experiments in limnocorrals showed that T. crassus has an advantage over C. vicinus when fish predation is high.Department of Biology III, University of Ulm     

Survival and development of five species of cyclopoid copepods in relation to food supply: experiments with algal food in a flow-through system

1. Cyclops spp. generally develop and grow during favourable food conditions in spring and undergo a diapause in summer, while Acanthocyclops robustus, Mesocyclops leuckarti and Thermocyclops crassus develop and grow in summer when they face poorer food conditions and more competition from Cladocera. Since nauplii are the bottleneck in copepod development, we tested the hypothesis that Cyclops abyssorum and C. vicinus nauplii have higher food requirements for survival and development than the nauplii of A. robustus, M. leuckarti and T. crassus. We also tested survivorship and development from hatching to adulthood. 2. Survivorship and development of the copepods was studied in a flow‐through system using five concentrations of the phytoflagellate Chlamydomonas reinhardtii in the range from 1 × 10⁴ to 4.5 × 10⁵ cells mL^?1 (approximately 0.5–22.5 mg C L^?1). 3. Nauplii of both species of Cyclops died at intermediate to low (C. abyssorum) and low (C. vicinus) food concentrations, while nauplii of A. robustus, M. leuckarti and T. crassus survived at all concentrations. 4. The negative effects of low food concentration were also reflected in development. In C. abyssorum and C. vicinus, development duration increased at low food concentration while development was much less affected in A. robustus and T. crassus. Mesocyclops leuckarti was intermediate between Cyclops spp. and A. robustus/T. crassus, with an increase in development duration at the lowest food concentration. 5. Our results support the hypothesis that summer diapause in Cyclops spp. has developed as a strategy to avoid a food bottleneck for nauplii.     

Filtering and feeding rates of cyclopoid copepods feeding on phytoplankton

The algal biomass ingested by omnivorous cyclopoid copepods (Cyclops kolensis and C. vicinus) was measured by two methods in the hypertrophic Heiligensee in Berlin (West Germany). The clearance and ingestion rates inferred from measurements of natural populations of ¹⁴C labelled phytoplankton were compared with those obtained from chlorophyll a determinations using the presence/absence method (observed chlorophyll a content of natural lake phytoplankton with and without addition of cyclopoids). Both methods gave similar results. Nevertheless, the radio tracer method is preferred, mainly because the short feeding duration excludes high variations in both the food composition and food concentration that limit the presence/absence method.     

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.     

Reproduction and adult longevity of five species of planktonic cyclopoid copepods reared on different diets: a comparative study

1. The nutritional value of a pure algal food, the phytoflagellate Chlamydomonas reinhardtii, and a mixed diet was tested for five planktonic cyclopoid copepods (Acanthocyclops robustus, Cyclops vicinus, Cyclops abyssorum, Mesocyclops leuckarti and Thermocyclops crassus). The algae were offered at high density (4.5 × 10⁵ cells ml^–1; 22.5 mg C l^–1) in a flow-through system. The mixed diet consisted of concentrated natural plankton (rotifers, copepod nauplii, small copepodites and large phytoplankton forms) in the size range 50–150 μm and with a dry mass > 20 mg l^–1. Reproductive parameters (clutch size, interclutch periods, number of clutches produced) and adult longevity were monitored as indicators of nutritive value. 2. All species had a significantly lower reproductive output and a shorter or unchanged adult lifespan on the algal compared with the mixed diet. 3. The species differed considerably in their ability to use algae. Mesocyclops leuckarti produced no clutches with algae, and females died earlier than with the mixed food. Acanthocyclops robustus and C. vicinus produced smaller and fewer clutches, displayed a longer interclutch period and shorter (A. robustus) or similar (C. vicinus) lifespan on the algal food than on the mixed food. Thermocyclops crassus and C. abyssorum produced smaller clutches with the algal food, but interclutch period was shorter with the algal than with the mixed diet (T. crassus) or of the same length with both diets (C. abyssorum). Adult lifespan was the same under both food regimes. Unfed females produced no eggs. 4. The ability to utilize algae, the reproductive output and the reproductive allocation were not related to body size. Acanthocyclops robustus, a species of intermediate size, produced by far the largest clutches and the most eggs per lifetime under both food regimes and invested more in reproduction than the other species. 5. Reproduction was costly. Unfed and non-reproducing females of C. vicinus and C. abyssorum reached the same age or lived longer than fed and reproducing ones. 6. The adult cyclopoids tested are primarily omnivorous, but utilize algae which are frequently sufficient for egg production.     

Life cycles of two limnetic cyclopoid copepods, Cyclops vicinus and Thermocyclops crassus, in two different habitats

The life cycles of Cyclops vicinus and Thermocyclops crassusin two shallow eutrophic habitats, Junsainuma and Naganuma Ponds,Hokkaido, Japan, were investigated. Both ponds exhibited similarseasonal patterns of temperature, oxygen levels and pH duringice-free periods; however, oxygen levels were extremely lowerunder the ice in Naganuma Pond. Cyclops vicinus showed differentlife cycles in the two ponds; in Junsainuma Pond, it reproducedin winter and spring (January-May) and entered diapause duringsummer and autumn (June-October) as copepodite IV stage, whileit reproduced in autumn (October-November) and spring (April-May),and entered diapause in summer (June-September) and winter (Januaryand February) as copepodite V stage in Naganuma Pond. Thermocyclopscrassus entered diapause during winter (December-April) as copepoditeIV and V stages in both ponds, and egg-bearing females appearedonly during the warmseason, from early May to late October,when water temperatures were >10°C. Summer diapause inC.vicinus was suggested to be an adaptation against fish predation,whereas C.vicinus entered winter diapause in Naganuma Pond probablyto avoid low oxygen levels. Thermocyclops crassus entered diapausein both ponds to avoid low water temperature. These resultssuggest that biotic and abiotic factors are important for leadingto specific life cycles of cyclopoid copepods in small waterbodies.     

Zooplankton in Loch Lomond: perspectives,predation and powan

Published and unpublished accounts of the zooplankton community of Loch Lomond are reviewed. Loch Lomond's zooplankton community is unique amongst those of Scotland's other large lochs. Holopedium gibberum appeared in Loch Lomond in the years between the studies of Slack & Hamilton in the 1950s and zooplankton surveys in the late 1970s.The annual cycles of Eudiaptomus gracilis, Cyclops strenuus abyssorum and Mesocyclops leuckarti are described for the mid basin of Loch Lomond. E. gracilis overwinter mainly as adult with variable egg production during their lifespan of 5–8 months. The cyclopoid copepods have variable life history characteristics. C. str. abyssorum produce a single generation per year, overwinter by means of resting eggs and a small population of planktonic adults, while M. leuckarti is present in the plankton throughout the year, but in low numbers during winter.Zooplankton populations in Loch Lomond are predated by the diverse fish community which includes a facultative planktivore in the powan, Coregonus lavaretus. Powan feed heavily on zooplankton from late spring until late autumn. Powan display selectivity in the prey species taken, in the size distribution and morph types taken within prey species. Zooplankton species, morphs and forms in Loch Lomond tend to reflect this predation.Zoology Department, University of St. AndrewsZoology Department, University of St. Andrews     

Coexistence with similar life cycles in two species of freshwater copepods (Crustacea)

Two species of freshwater cyclopoid copepods, Cyclops scutifer Sars and Mesocyclops leuckarti (Claus), with different ecological requirements, coexist in three humic ponds near the timberline in boreal forests in south-central Norway. The life cycle in both species is basically regulated by complete depletion of oxygen during the long winter. There is a short period of reproduction in spring followed by a rapid development during summer and autumn to advanced instars, capable of diapause when oxygen depletion starts in early winter. Both species have an exclusive 1-yr cycle.The long winter under anoxic conditions is passed in diapause as large copepodids (late developmental instars) with usually no individuals left in the plankton.Even if co-occurring in the same habitats with similar life cycles, there is a temporal and spatial segregation.     

Coexistence of the predatory cyclopoids Acanthocyclops robustus (Sars) and Mesocyclops leuckarti (Claus) in a small eutrophic lake

The coexistence of two common species of predatory cyclopoid copepods, Acanthocyclops robustus and Mesocyclops leuckarti, in a small, eutrophic lake was studied. Both species are of similar size, inhabit the same water layers and have similar seasonal occurrence. A. robustus shows faster embryonic and postembryonic development and produces bigger clutches. Ingestion rates of A. robustus are higher than in M. leuckarti, especially with large prey types and low temperatures. Females of A. robustus are heavily preyed upon by fish. This predation pressure leads to extremely skewed sex ratios, which are unfavourable to population growth. A. robustus is therefore, on balance, favoured in productive lakes. Dominance over M. leuckarti is, however, less clear in warmer years.     

A simple method for maintaining a laboratory supply of limnetic cyclopoid copepods

The following method is used for maintaining a laboratory supply of freshwater cyclopoid copepods with intervening diapause phase. Animals are collected with a core sampler from the upper sediment surface just after initiation of diapause, i.e. during early summer for summer resting species and mid autumn for winter resting species. The method makes it possible at all times of the year to keep a variety of cyclopoid copepod species ready for experimental work. The following widespread species have been used: Mesocyclops leuckarti, Thermocyclops oithonoides, Diacyclops bicuspidatus, Cyclops vicinus, C. strenuus and C. abyssorum. Probably all cyclopoid copepods with a diapause phase could be stored according to the present method.     

The life cycle of Cyclops vicinus in Lake Søbygård: new aspects derived from sediment analyses

Cyclops vicinus is the only copepod species in the pelagic zone of Lake Søbygård and can be found there all year round. We studied the population dynamics of this copepod over a one-year period. In contrast to earlier studies we included the copepods in the pelagic zone as well as the copepods resting in the sediment. Cyclops vicinuswas found not only in the open water, but also in the sediment throughout the year. From the fluctuations of the abundances in both habitats we suggest that the life cycle is more complex than known so far: different diapause pattern appear within the population and the induction of diapause is different than in other populations. We assume that these features contribute to the predominance of C. vicinusin Lake Søbygård.     

Nutritional suitability of the dinoflagellate Ceratium furcoides for four copepod species

The nutritional suitability of Ceratium furcoides for Eudiaplomusgracilis, Mesocyclops leuckarti, Thermocyclops oithonoides andCyclops abyssorum was studied by observing ingestion and assimilation.The dinoflagellate was ingested by female adults of all species.The calanoid copepod E.gracilis could not utilize Ceratium;mortality was high and no egg production was observed when Ceratiumwas the only food source. Mortality was low for the adult cyclopoidson this food and reproduction indicated that Ceratium was assimilated.The first two copepodite instars of M.leuckarti were not ableto handle Ceratium, while older stages preyed on them. The dinoflagellatewas not ingested by female M.leuckarti when its densities werelow. Advanced copepodite stages of C.abyssorum developed intoadults on a diet of Ceratium only. Mesocyclops leuckarti femalesingested Ceratium when offered a mixed food source of Ceratiumand the rotifer Brachionus rubens, but the rotifer was positivelyselected, even if its density was low. The results show thatCeratium provides a suitable food source for advanced copepoditeinstars and adult cyclopoid copepods, although it is not a preferredfood source.     

Comparative feeding ecology of Tropocyclops prasinus mexicanus (Copepoda, Cyclopoida)

The herbivorous and carnivorous feeding biology of Tropocyclopsprasinus mexicanus, an especially small cyclopoid copepod, wasstudied under in situ food conditions in three different lakesand under experimentally augmented food supplies. The mass-specificfood uptake is compared to that of two larger species—Cyclopskolensis and Cyclops vicinus. Under in situ food conditions,T.p.mexicanus depended to a larger extent on algae than invertebrateprey and showed lower mass-specific ingestion rates than thetwo larger species. Daily mass-specific uptake rates for algaeranged between 10 and 24% of its body mass versus 0.7–7%for invertebrate prey. The larger species C.vicinus and C.kolensisingested a total dry mass equivalent to 106% or up to 143% oftheir body mass with algae contributing 66 and 81%. However,under enriched food availability, T.p.mexicanus is able to ingesta biomass equivalent to its body mass, with an algae (54%) andprey (40%) portion similar to that of the larger species. Bodysize appears to be an important factor for the relative importanceof algal versus invertebrate prey for cyclopoid copepods.