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.     

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.     

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.     

Herbivorous nutrition of Cyclops vicinus: the effect of a pure algal diet on feeding, development, reproduction and life cycle

Nutritional requirements, functional response, development andreproduction of Cyclops vicinus were studied with exclusivelyalgal food. Phytoflagellates were found to be adequate foodresources for both juvenile development and egg production.Ingestion measurements were performed with Chlamydomonas reinhardii.The functional response data give evidence for low feeding efficiency,especially for the naupliar stages. A difference between naupliiand older instars was also found in their quantitative foodneeds. A higher threshold food concentration was observed fornaupliar development (0.4 mg C l^–1) than for copepoditedevelopment (0.2 mg C l^–1). The calculation of assimilationefficiencies suggests that the high food requirements are dueto low specific ingestion rates rather than poor assimilationefficiency. Development time decreased as algal density increasedand males developed more quickly than females at all food concentrations.Body size and carbon content increased with increasing foodconcentration. Continuous egg production was observed abovea food concentration of 0.5 mg C l^–1. The results haveimplications for the life cycle of C. vicinus. Summer diapauseis interpreted as a strategy to avoid starvation of the juvenilestages. The facultative herbivory of the adults might be anadvantage when competing against other more carnivorous cyclopoidcopepods. ¹Present address: Agricultural University of Wageningen, Departmentof Mathematics, Dreijenlaan 4, NL-6703 HA Wageningen, The Netherlands     

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.     

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.     

Remote detection of algae by copepods: responses to algal size, odors and motility

Selective feeding on large algae by copepods involves remotedetection of individual particles and subsequent active captureresponses In this study we use radiotracer experiments to quantifythe clearance rates of five coexisting freshwater copepods andto investigate the relative merits of the chemoreception andmechanoreception hypotheses of remote detection Tropocyclopsand three diaptomid copepods exhibited relatively high clearancerates when feeding on low concentrations of large algae, suggestingthat most previous studies with freshwater copepods have underestimatedmaximal clearance rates and the degree of size selectivity.All five species of copepods exihibited strong selection foran intermediate-sized flagellate (25 µm Cartena) or alarge-sized nonmotile alga (80 µ.m Pediastrum) over asmall-sized flagellate (6 µ.m Chlamydomonas). The weight-specificclearance rate for Tropocylops prasmus feeding on motile Cartena(271 ml mg^–1 h^–1) was about twice that of threediaptomid copepods and more than an order-of-magnitude higherthan the estimate for Epischura lacustris feeding on its preferredalga, Pediastrum Assuming that distance chemoreception (‘smell’)is important in remote detection, we predicted that the additionof high concentrations of ‘algal odors’ would obscureany chemical gradients emanating from individual algal cellsand would thereby hinder the remote detection and active captureof large algae Contrary to this hypothesis, the addition ofamino acids, sucrose, and algal extracts had no effect on theclearance rates and selectivity of Diaptomus birgei. These results,together with recent cinematographic studies (Vanderploeg etal.,1990), suggest that mechanoreception is the primary mechanismfor the remote detection of large particles by diaptomid copepods.A raptorial cyclopoid, Tropocyclops prasinus, exhibited strongpreferences for motile algae, whereas a suspension-feeding calanoid,D birgei, did not select between motile and nonmotile cells.Motility appears to be an important factor in algal detectionfor small cyclopoid copepods but not for suspension-feedingdiaptomids     

Omnivory in cyclopoid copepods: comparisons of algae and invertebrates as food for three, differenfly sized species

We examined the importance of algal versus invertebrate preyfor three cyclopoid copepods by comparing reproductive successfor females fed on three diets: exclusively algal food, exclusivelyinvertebrate prey and a combination of algal plus invertebrateprey. The three cyclopoid species represent a distinct gradientin body size: Tropocyclops prasinus mexicanus (small body size),Diacyclops thomasi (medium body size) and Mesocyclops edax (largebody size). Our results provide strong evidence for the potentialimportance of algae in the diet of adult cyclopoid copepods.However, the role of herbivory is dependent on species bodysize. We found that algae are the key dietary factor for reproductivesuccess of the small T.p.mexicanus. In contrast, the availabilityof invertebrate prey had a greater influence on reproductionby the two larger species. Overall, cyclopoid copepods shouldbe generally considered as having the potential to derive theirenergy from either animal or algal sources. This has importantconsequences for interpretations of the forces influencing zooplanktoncommunity diversity as well as for general food web theory.     

Life history, biomass and production of two planktonic cyclopoid copepods in a shallow subtropical reservoir

Two planktonic cyclopoid copepods (Tropocyclops prasinus andMesocyclops longisetus) were raised in the laboratory to obtainlife history information (duration of embryonic and post-embryonicdevelopment, reproductive performance, longevity, and stage-specificlength and weight values). Animals were maintained at 20 and25°C, and fed ad libitum. Development times were temperaturedependent when food was not limiting, with shorter periods ofembryonic and post-embryonic development and decreased longevityat 25°C. Laboratory data on the duration of developmentand biomass, together with population dynamics data obtainedin the field, were used to estimate summer and winter biomassand production of these species in a shallow reservoir, LagoaDourada, Brazil. The maximum production rate of T. prasinus,attained during summer, was 2.8 mg dry weight (DW) m^–3day^–1 and the highest daily production:biomass (P:B) ratiowas 0.29, whereas for M. longisetus the maximum production ratewas 1.4 mg DW m^–3 day^–1 and the highest daily P:Bratio was 0.39, in the winter. Over short time intervals (everyother day), there was great variability of the species productionrates. Species production rates were low compared to valuesreported in the literature for the same or other species ofequivalent sized copepods from both tropical and subtropicalregions.     

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.     

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.     

Life cycle of Cyclops vicinus in relation to food availability, predation, diapause and temperature

The seasonal cycle of Cyclops vicinus was studied during a 5year period in the shallow and hypertrophic Lake Søbygård.The annual number of generations varied between three and sixand sometimes included midsummer generations. Naupliar and copepoditedensity was extremely high, the maximum recorded being 1313and 745 1^–1 respectively. The seasonal fluctuation inthe abundance of C.vicinus was related to the concentrationof edible phytoplankton (EDP): when diatoms and cryptomonadswere abundant, nauplii and copepodites were also abundant, whereaswhen greens such as Scenedesmus spp. were dominant, no cohortswere observed. The maximum density of copepodites in the separatecohorts and the amount of EDP, expressed as carbon content,were found to be significantly correlated. The development ofthe second cohort also seemed to depend on the occurrence ofa peak in rotifer density. High number of eggs per female generallycoincided with high biovolume of EDP, suggesting that algaeconstituted a major part of the diet of adult C. vicinus. Theduration of the naupliar stage was inversely related to temperatureand is one of the shortest times that have been recorded inthe literature. The short duration is probably attributableto the abundance of food. Fish composition and the calculatedsex ratio of C.vicinus indicate low predation pressure fromplanktivorous fish. The high density of C.vicinus during thesummer, when the whole population would be expected to haveentered diapause, is probably attributable to the concurrenceof low predation and the abundance of food.     

Diel periodicity and selectivity in the feeding rate of the predatory copepod Mesocyclops edax

The diel periodicity and selectivity in the feeding behaviorof the predatory cyclopoid copepod Mesocyclops edax was investigatedat 3 h intervals over two 24 h sampling periods in nature. Gutcontent analyses revealed an increase in gut fullness at 11.30and 20.30 in July, and at 20.30 in August. The increase in gutfullness at these times could not be explained by an increasein prey density, changes in predator-prey overlap or differencesin prey vulnerability. We suggest that M.edax exhibits a truediel periodicity in the intensity of its predatory activities,although the alternative hypothesis of a diel periodicity ingut passage time cannot be ruled out. Vanderploeg and Scavia'sE^* selectivity index indicated a preference for rotifer andcladoceran prey over copepods.     

Response of ciliates and Cryptomonas to the spring cohort of a cyclopoid copepod in a shallow hypereutrophic lake

The impact of a cyclopoid copepod population on the protozoacommunity (two ciliate categories and Cryptomonas) was assessedweekly during the spring cohort of Cyclops vicinus (one monthduration) in hypereutrophic Lake Søbygård by insitu gradient experiments with manipulation of ambient zooplanktonabundance. As C.vicinus always made up >92% of the zooplanktonbiomass, the response of protozoa is assumed to be a resultof predation by the copepod. Significant effects of copepodbiomass on protozoa net population growth rates were obtainedin the four experiments. Copepod clearance rates were significantlyhigher on oligotrichs than on prostomatids and Cryptomonas butdeclined for all three protozoa categories during the firstthree weeks of the copepod cohort, probably because of the changein developmental instar composition of the copepod population.Grazing impact on protozoa at ambient copepod abundance wasconsiderable (range, 0.05–0.87 day^–1) and could,together with the estimated reproductive potential of protozoans(range, –0.20–0.87 day^–1), account for thedecline in abundance and biomass of protozoa during the cohortdevelopment. Carbon flow from the protozoa to C.vicinus (range,2.8–23.5 µg C l^–1 day^–1) documents thepresence of a trophic link between protozoa and the spring cohortof C.vicinus in Lake Søbygård.     

Mating frequency and interspecific matings in some freshwater cyclopoid copepods

Laboratory investigations with Cyclops vicinus and Metacyclops minutus documented that mating frequency decreased significantly after the initial mating, i.e. re-mating is relatively rare in cyclopoid copepods. Re-mating is unnecessary because females are able to fertilize multiple clutches of viable eggs from one insemination. Similar sized Cyclops vicinus and Cyclops furcifer interbreed frequently. Interbreeding does not occur when size differences and taxonomic differences are as great as between C. vicinus and M. leuckarti. In C. vicinus and M. leuckarti, the duration of the last mating phase (spermatophore transfer until release of the female) and consequently the duration of the entire mating process, is different. I conclude that the low re-mating frequency in cyclopoid copepods probably evolved to accelerate clutch production in unpredictable environments and to reduce predation risk. The occurrence of interspecific mating discourages the co-existence of similarly sized, related species. Mating behaviour may have a considerable influence on zooplankton community structure.     

The production and ingestion of faecal pellets by nauplii of marine calanoid copepods

The downward transport of organic matter as zooplankton faecalmaterial is influenced by copepods which fragment, ingest andrecycle some of the pellet contents. Most of this activity hasbeen attributed to the later copepodite stages and the adults,but little is known about the role of nauplii. Stage-relateddefaecation rates during the naupliar development of two speciesof copepod, Calanus helgolandicus and Pseudocalanus elongatus,were quantified in a series of laboratory experiments. The productionof faecal material commenced soon after the appearance of theNIII in both species and increased throughout naupliar development.The causes of the increase were the formation of larger pelletsby later stages in Calanus and an increased rate of productionby Pseudocalanus. Calanus nauplii, when supplied with algalfood at concentrations that would support full naupliar development,ingested or broke up the pellets of the smaller Pseudocalanusspecies at rates of 1.15 pellets nauplius^–1 h^–1This consumption increased to 2.96 pellets nauplius^–1h^–1 when the concentration of algal food was reduced toa limiting level. Pseudocalanus was not able to consume thepellets of Calanus. Ingestion of Pseudocalanus faecal pelletsby Calanus could supply a nutritional benefit to a food-limitednauplius.     

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.     

Steady-state rotifer growth in a two-stage, computer-controlled turbidostat

Steady-state populations of rotifers (Brachionus calyciflorus)were maintained in twostage, continuous-flow turbidostatic cultureon the green alga Chlorella pyrenoidosa. In this system, themaximum specific growth rate,µ^max of the rotifers wasmaintained by using a computer to control the concentrationof algae, as rotifer food, in the rotifer culture. As rotifersconsumed algae, the turbidity decreased until a set-point wasreached. Then fresh algal suspension (supplied from a steady-statealgal chemostat) was metered into the rotifer culture, whichwas held in the dark. Rotifer and algal populations, as wellas rotifer µ^max entered steady states. These steady-stateresults were consistent with previous data from chemostat studies,but growth transients indicated that the of the µ^maxrotifersmay be subject to selection. The system is unique in providinga means to explore population dynamics of a metazoan maintainednear its µ^max.     

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.     

Effects of temperature, salinity and food level on the life history traits of the marine rotifer Synchaera cecilia valentina, n. subsp.

A strain of the marine rotifer Synchaeta cecilia valentina,n. subsp., isolated from the Hondo of Elche Spanish Mediterraneancoastal lagoon at 22 salinity, was cultured in the laboratoryin 20 ml test tubes and fed with the alga Tetrasemis suecica.The effect of two temperatures (20 and 24°C), four salinities(20,25,30 and 37) and two food levels (15 000 and 25000 cellsml^–1) on the life history traits of this rotifer werestudied in life tables performed with replicated individualcultures. Temperature and salinity had a significant negativeeffect (P < 0.001) on the average lifespan (LS) and on thenumber of offspring per female (R₀) The effect of food levelon LS is unclear, whereas R₀ is greater at 20°C with thelower concentration of algae and at 24°C with the higheralgal concentration. The maximum values of LS and R₀, 5.6 daysand 9.2 offspring per female, respectively, were recorded at20°C, 25o salinity and low food concentration. There isalso a clear negative effect on the intrinsic growth rate (r)due to salinity. The effect of temperature depends on the foodlevel and, as occurs with R₀ the maximum values of r occur withthe lower algal concentration at 20°C, whereas at 24°Cthey are obtained with the higher algal concentration. Theser values, from 1.04 to 1.10 day^–1, were reached at 24°C,salinities of 20–25 and with high food concentration.