The role of photoreception in the swarming behavior of the copepod Dioithona oculata

The copepod Dioithona oculata forms dense swarms near mangrove prop roots that are centered around shafts of light penetrating the mangrove canopy. Swarms can be created in the laboratory within light shafts created with a fiber optic light pipe. Laboratory observations of swarming behavior were recorded using video cameras, and the swimming behavior of the copepods and density of the swarms were quantified using video‐computer motion and image analysis techniques. Swarm formation results from a combination of phototactic and klino‐kinetic behavior. Dark adapted copepods initially exhibit a photophobic response to a light shaft, but become positively phototactic within 3–5 min after exposure to the light. Copepod aggregation rates under the light fit a saturation model, suggesting that copepods are attracted independently to the swarm marker. Copepods reverse their swimming direction when they encounter light intensity gradients near the edge of a light shaft, which aids in maintaining the swarm. Swarm formation can occur in the laboratory at light intensities as slow as 0.1 μM photons m^‐2 s^‐1, which is similar to light intensities at dawn when they are first observed to form in nature. Swarm formation appears to have an endogenous rhythm, as copepods will not form swarms at night under a light shaft.     

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.     

Behavioral interactions between a cyclopoid copepod predator and its prey

Behavioral observations on the predatory interactions betweenMesocyclops edax and several different types and sizes of preyrevealed that prey size alone was less important than otherspecific morphological and behavioral characteristics of theprey in deterring successful predation by the copepod. The behavioralresponses of Bosmina and Asplanchna to an attacking copepodwere passive and consisted of a simple retraction of vulnerableswimming appendages which made the prey more difficult to grasp.Daphnia and Diaphanosoma on the other hand exhibited very activeswimming escape responses. Tropocyclops usually avoided M. edaxby fleeing before the larger predator could detect them. Thehard carapaces of Daphnia, Bosmina and Keratella were effectiveat reducing ingestion following capture by M. edax. The resultsof these behavioral observations were supported by enclosureexperiments in which the predator was offered a choice betweentwo prey simultaneously. Cyclopoid copepods are capable of successfullyattacking, capturing and ingesting prey organisms several timestheir own body length. Although size alone may influence thepreference of cyclopoid copepods on large and small individualsof the same or similar prey species, it is not a dependabledeterminant of the preference of cyclopoids on multispecificprey assemblages. ¹Present address: Department of Biology, Williams Hall No. 31,Lehigh University, Bethlehem, PA 18015, USA     

Egg production, growth and development of the cyclopoid copepod Oithona similis

Egg production, growth and development rates of Oithona similiswere measured in the laboratory as a function of food concentrationand composition. On an optimum diet, development is isochronaland growth is near exponential. The maximum juvenile growthrate at 15C (0.2 day^–1) is similar to juvenile growthin calanoid copepods. The maximum weight-specific egg productionrate (0.1 day^–1), on the other hand, is substantiallyless than in free-spawning calanoids, but similar to that inegg-carrying calanoids. In the Kattegat, Oithona spp. egg productionis strongly limited by food during summer and controlled bytemperature during winter. The seasonal signal in fecundityand population biomass is much weaker than in the co-occurringfree-spawning calanoid genera, where fecundity and populationbiomass undergo dramatic seasonal viaration. ¹Present address: Instituto Nacional de Investigacin y DesarrolloPesquero, INIDEP, CC 175, Playa Grande, Mar del Plata, Argentina     

Laboratory and field experiments on the feeding ecology of the cyclopoid copepod, Mesocyclops edax

SUMMARY. 1. Gut contents of Mesocyclops edax consisted of rotifers. cladocerans, copepods and algae, the importance of each varying seasonally.
2. Functional response experiments using two soft-bodied rotifers as prey showed an increase in ingestion rate with increasing prey density up to a threshold, followed by a decline.
3. Enclosure experiments with natural assemblages of prey from two lakes indicated that Mesocyclops edax is a selective feeder and may ingest fifty or more prey per predator per day, accounting for up to 17% of the standing crop of its preferred prey species per day in nature.     

Variable life history of a cyclopoid copepod: the role of food availability

Populations of the copepod Cyclops vicinus in two Danish lakes differed in their life cycles. In Lake Væng C. vicinus was absent during summer, whereas in Lake Søbygård it continued producing distinct cohorts throughout summer. A comparison between the lakes showed that in the lake where C. vicinus was absent during summer, food limitation generally existed. In contrast, abundance of C. vicinus during summer in Lake Søbygård always coincided with available resources sufficient for the whole life cycle. Thus, summer diapause of C. vicinus is suggested to be a strategy to avoid food limitation. Avoidance of fish predation does not seem substantially to influence life cycles.     

Changes in body-length, and in shape of the furcal ramus, of a freshwater cyclopoid copepod

Some morphological relationships during larval development of a freshwater cyclopoid copepod (Crustacea), Cyclops strenuus abyssorum Sars (Gurney, 1933) have been studied. The proportion which the cephalothorax forms of total length decreases at each copepodid moult whilst that which the furcal ramus forms increases. However, within each instar, each of these ratios is a constant at different seasons and in different lakes. The amount by which total length in one instar is greater than that in the preceding instar tends to decrease at each moult from a maximum of 1.31 in the first copepodid moult to lower values, especially in males. The furcal ramus is twice as long as wide in the first copepodid instar but seven times as long as wide in the adult female. This change is enhanced in the last two moults not only by a proportionately greater increase in length over width but by an absolute reduction in width itself.     

Planktonic predators and copepod abundance near the Dutch coast

Hypotheses that planktonic predators are responsible for thespring-summer decrease in copepod abundance and that the dominantpredator, Pleurobrachia pileus, is associated with high concentrationsof copepods were investigated at a station near the Dutch coast.Neither hypothesis was supported. Predators and copepods weresampled together with a 156 L ‘water box’ from lateApril through early July, including the season of P.pileus abundance.Using predators and copepods from the same box samples, hencefrom the same water parcel, feeding rates on copepod naupliiand copepodites + adults were measured onboard ship. Less than6% of the copepods, the sensitivity of the method, were removedper day. In additional shipboard feeding experiments net-caughtP.pileus were added to ambient copepod densities. By combiningwater volume cleared of copepods with ambient P.pileus densityin the sea, the predicted impact was 0–1.6% of copepodseaten per day from late April to early July. The hypothesisthat P.pileus associates with copepod concentrations was testedby comparing abundances of both groups from the same box samples.The correlations were not significant for any sample series.The ways measurement methods have restricted progress in understandingthe predatory impact of Pleurobrachia sp. are also considered.     

Food limitation in a wild cyclopoid copepod population: direct and indirect life history responses

We studied the effects of food limitation on the population dynamics of the freshwater cyclopoid copepod Diacyclops thomasi in Oneida Lake, New York. In the field population, maximum juvenile abundance coincided seasonally with high phytoflagellate concentration. During the clear-water phase (a seasonal period of low algal density), D. thomasi disappeared from the water column, but fourth-instar copepodids (CIV) were found encysted in developmental arrest in the sediment. Laboratory assays of the effect of the density of two types of food on copepod life history parameters showed that temporal variation in the concentration of relatively small phytoflagellates significantly affected stage-specific development times. This food limitation was most pronounced during the clear-water phase, and supplementation of the diet with a laboratory-cultured phytoflagellate, Chlamydomonas, prevented food limitation. Although developmental arrest appears to be controlled primarily by photoperiod, availability of the larger, more mobile food, Euglena, also influenced the percentage of individuals entering developmental arrest in the laboratory. An investigation of the spatial and temporal emergence pattern in the field revealed that CIV copepodids started to emerge in late autumn and that emergence rates were significantly greater at deep-water locations (9–12 m water depth) compared with shallow-water locations (5–7 m). The clear-water phase in Oneida Lake is an annual event, probably produced by intense grazing by Daphnia pulicaria and Daphnia galeata. Food limitation is thus very likely a recurrent phenomenon for D. thomasi. This apparent seasonal competitive impact of Daphnia on Diacyclops affects both nauplii and immature copepodids. Diacyclops shows two types of responses to the food limitation: (1) the physiological response of slowed active development, and (2) the adaptive response of developmental arrest. Received: 3 November 1997 / Accepted: 1 March 1998     

A new marine cyclopoid copepod of the genus Neocyclops (Cyclopidae,Halicyclopinae) from Korea

A new cyclopoid species of the genus Neocyclops Gurney, 1927 is described. Type specimens were collected from a beach on south-western coast of the Korean Peninsula by rinsing intertidal coarse sandy sediments. Neocyclops hoonsooi sp. n. is most characteristic in showing the conspicuous chitinized transverse ridges originating from the medial margins of the coxae of all swimming legs. The new species is most similar to Neocyclops vicinus, described from the Brazilian coast, and Neocyclops petkovskii, from Australia. All three species share a large body size (more than 750 µm long), the presence of an exopodal seta on the antenna, two setae on the mandibular palp, the same seta/spine armature on the third endopodal segment of leg 3 (3 setae + 3 spines), and the fairly long inner distal spine on the third endopodal segment of the female leg 4. However, Neocyclops hoonsooi sp. n. differs from both species by the much shorter caudal rami (less than 1.7 times as long as wide) and the shorter dorsal caudal seta VII. Furthermore, Neocyclops hoonsooi is clearly distinguished from Neocyclops vicinus by the 10-segmented antennule (vs 12 segments in Neocyclops vicinus), and from Neocyclops petkovskii by the elongate inner distal spine on leg 5 exopod and the 3-segmented leg 5 in male (vs 4-segmented in Neocyclops petkovskii). A tabular comparison of characters separating Neocyclops hoonsooi from its closest allies and a key to Neocyclops species from the Indo-Pacific Ocean are provided. This is the first record of the genus Neocyclops from the northern Pacific.     

Predation by juvenile roach on the calanoid copepod Eudiaptomus gracilis and the cyclopoid copepod Cyclops vicinus: a laboratory investigation with mixed and single prey

We tested the outcome of predation by juvenile roach on the calanoid copepod Eudiaptomus gracilis and the similar sized cyclopoid copepod Cyclops vicinus in laboratory experiments with mixed and single-prey. When ovigerous females and adult non-ovigerous females of the calanoid copepod and the cyclopoid copepod were offered in equal numbers to the fish in the mixed-prey experiments, the cyclopoid copepods were significantly more heavily preyed upon than the calanoids. Between 14 and 16 females of the cyclopoid copepod and only between 2 and 4 calanoids had been consumed after the observation period of two hours. The single-prey experiments revealed that the conspicuousness of the calanoid vs the cyclopoid depended on sex and gravidity. Ovigerous females of the cyclopoid copepod were slightly earlier detected by the fish than ovigerous females of the calanoid, probably a result of the highly visible egg-sacs carried by the cyclopoid females. Females without eggs and males of the cyclopoid copepod were recognized later by the predator than females without eggs or males of the calanoid, probably a result of the different behaviours of the different copepod taxa. Cyclopoids frequently congregated near the aquarium bottom while calanoids were closer to the water surface where they were better visible. The calanoid copepod could better escape the fish's attacks than the cyclopoid copepod. Egg-bearing females of both the calanoid and the cyclopoid copepod could significantly better escape than non-ovigerous females or males. Probably ovigerous females react very early to water disturbances caused by the predator. Activity measurements showed that the cyclopoid copepod displayed 2 to 4 times more hops per time unit than the calanoid. Probably the high number of jerky movements displayed by the cyclopoid attracted attention of the predator and contributed to its greater vulnerability.     

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.     

Post-embryonic developmental rates as a function of food type in the cyclopoid copepod, Mesocyclops thermocyclopoides Harada

We studied under laboratory conditions the post-embryonic developmentalrates (from nauplii to adult stage) of Mesocyclops thermocyclopoides,commonly found in the eutrophic and hypertrophic local ponds,in relation to different food types. Nauplii hatched from theeggs collected from laboratory-maintained mass cultures werereared at 25l.5C, using the following test diets: bacteria(Klebsiella aerogenes), cyanobacteria (Microcystis aeruginosa,as single cells and in the form of colonies), green algae (Chlorella,Scenedesmus and Chlorogonium), ciliates (Tetrahymena) and rotifers(Brachionus angularis). The developmental rates were fastest,and the proportion of nauplii reaching the copepodid stage andthat of copepodids reaching the adult stage the highest, withChlorogonium, followed by ciliates and rotifers. Developmentwas incomplete with bacteria and Microcys-tis.Ttie algae Chlorellaand Scenedesmus supported complete post-embryonic developmentonly when cultured in media enriched with beef extract, bactotryptoneand yeast extract. With rotifers as the exclusive food, >70%of the nauplii reached the copepodid stage, and >85% of thecopepodids the adult stage. The ratio (D_c/D_n) of copepodid duration(D_c) and naupliar duration (D_n) was significantly differentfrom 1.0, indicating the absence of isochronal development inM.thermocyclopoides. Our results show that post-embryonic developmentin this species is not possible with certain algal diets, probablybecause of nutritional deficiencies, and that the later stagenauplii are capable of capturing rotifers and utilizing themas food to complete their development to the adult stage.     

Population structure of the intertidal copepod Tigriopus californicus as revealed by field manipulation of allele frequencies

Summary Allele frequencies in natural T. californicus populations were perturbed by introduction of copepods from neighboring differentiated populations. In five experiments, the Gpt^F allele was introduced into single recipient pools at a frequency of approximately 20%. In each case, the introduced allele declined to low frequencies (<3%) in less than one month, apparently due to dilution by residents of other pools on the same outcrop. In a larger scale experiment, the Pgi^F was introduced into four pools on a single small rock outcrop; all pools on the outcrop were subsequently monitored. While the allele frequency fell from approximately 40% to 10% during the first six weeks after the transplant, no further change in frequency was observed for the duration of the experiment (16 months). Within six weeks some spread of the allele to non-recipient pools on the same outcrop was observed; by eight months, allele frequencies in all pools on the outcrop were similar. Hence, despite the extensive turnover of subpopulations as single pools evaporate or are washed out, genetic homogeneity and stability of entire outcrops are maintained via extensive inter-pool gene flow; this contrasts sharply with the highly restricted levels of inter-outcrop gene flow.     

Effect of culture density and antioxidants on naupliar production and gene expression of the cyclopoid copepod, Paracyclopina nana

Although attempts have been made to use mass cultures of marine copepods as live foods in marine aquaculture, some limitations such as low density culture still exist. The brackish water cyclopoid copepod, Paracyclopina nana has the potential for mass culturing as live food. In this study, we not only investigated the effect of culture density on the naupliar production and specific gene expressions of P. nana, but also the effect of several antioxidants under the conditions of a high density culture. The naupliar production of the copepod decreased with increasing culture density. The expression of glutathione reductase (GR), selenium-dependent glutathione peroxidase (SeGPx), glutathione S-transferase kappa (GST kappa), heat shock protein 40 (Hsp40), and Hsp70 genes of P. nana increased in the high density treatment but vitellogenin genes (Vg1 and Vg2) showed downregulation. In the condition with 20 inds./mL, vitamin C had a significant decrease but sodium selenite induced the naupliar production of P. nana greatly. The expressions of GR, SeGPx, Hsp70, and Vg genes increased with the vitamin C treatment. Sodium selenite caused a decrease of SeGPx and Hsp40 but GST kappa increased in the treatment with 20 inds./mL. These results suggest that sodium selenite is a positive antioxidant which can increase the culture efficiency of the copepod.     

A new interstitial cyclopoid copepod from a sandy beach on the western shore of Lake Baikal,Siberia

A new species of cyclopoid copepod, Diacyclops biceri sp. nov., is described from Lake Baikal. It was found in the interstitial water of a sandy beach at Buchta Peschanaya on the western shore of the central basin of Baikal. The new species is unique in possessing 2-segmented endopods in swimming legs 3 and 4. Swimming legs 2 to 4 have 3-segmented exopods. The slender body form, the lack of the antennal exopodal seta, and the presence of a secondary pseudosomite anterior to the genital double somite of the adult female are interpreted as adaptations to the interstitial habitat. The harpacticoid Epactophanes richardi Mrazek was found in the same interstitial habitat as D. biceri.     

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.     

Epizootiology of Amblyospora connecticus (Microsporida) in field populations of the saltmarsh mosquito, Aedes cantator, and the cyclopoid copepod, Acanthocyclops vernalis

The natural ecology of a heterosporous microsporidium, Amblyospora connecticus was investigated at three different salt marsh habitats during 1986-1989. The parasite has a well-defined seasonal transmission cycle that occurs regularly each year and intimately involves the primary mosquito host, Aedes cantator, and the intermediate copepod host, Acanthocyclops vernalis. In the spring, the microsporidium is horizontally transmitted from the copepod, where it appears to overwinter, to the mosquito via the ingestion of haploid spores produced in the copepod. Mosquitoes develop a benign infection, and females transmit the microsporidium transovarially to their progeny via infected eggs. Oviposition occurs during the summer and infected eggs hatch synchronously in the fall causing widespread epizootics. Infected larvae die, and the cycle is completed when meiospores are released into the pool and subsequently are eaten by A. vernalis, which reappears in the fall and early winter. Amblyospora connecticus thereby persists by surviving in one of two living hosts throughout most of its life cycle rather than in the extra-corporeal environment. This represents an important survival strategy for A. connecticus as results show the salt marsh habitat to be a relatively unstable environment that is subject to periodic flooding and drying. The adaptive significance of utilizing an intermediate host in the life cycle is discussed as it directly facilitates transmission and enhances survival of the microsporidium.