The fine structure of a cephalic sensory receptor in the copepod Doropygus seclusus Illg (Crustacea: Copepoda: Notodelphyidae)

A cephalic organ of presumed sensory function is described in nauplii and copepodids of the ascidicolous copepod Doropygus seclusus Illg. The receptor, located bilaterally in the anterodorsal head region, is composed of dendrites of extra optic protocerebral origin which have ciliary protrusions with basal bodies, no rootlets, and a basal infrastructure of the 9 + 0 type. The cilia do not branch and their distal terminations contain only one to four microtubules. In nauplii and free-living copepodids, a large epidermal supporting cell encapsulates the end of one dendrite and its cilia in a sac. Other dendrites and their cilia pass through the supporting cell and, terminally, the cilia escape to form a whorled fascicle which contacts the anterolateral cephalic cuticle. The latter end organ reaches its greatest development in the second copepodid stage — the stage which infects the ascidian. All of the symbiotic stages of the copepod have only a proportionately smaller end organ of the saccular type and apparently lack the end organ consisting of whorls of ciliary ends. The function of the receptor is unknown, but it is suggested that the end organ which disappears in the symbiotic stages functions in second copepodids in host recognition.     

Direct evidence of tidally oriented behavior of the copepod <Emphasis Type="Italic">Eurytemora affinis</Emphasis> in the Seine estuary

The calanoid copepod Eurytemora affinis dominates the zooplankton communities in most northern hemisphere estuaries. A recurrent behavior noticed in several estuaries suggests that this species maintains its horizontal position through vertical migration. In order to investigate this behavioral strategy in detail, we sampled E. affinis nauplii, copepodids and adults at high frequency at a fixed point in the low salinity zone of the Seine estuary. We took water samples in sub-surface and near bottom during a 12- and 50-h tide cycle, in May 2002 and May 2004, respectively. Since nauplii have weak swimming capabilities compared to copepodid and adult stages, we proposed here to consider them as passive particles. The variation of the relative abundances of nauplii to older stages regarding the vertical position was therefore viewed as direct evidence of the behavior of later stages. We showed that copepodids and adults tried to stay close to the bottom during the ebb, were re-suspended in the water column 1 h after the low tide during the maximum current velocity, and then vertically migrated back to the bottom layer when current velocity decreased. We also showed that copepodids behaved as adults, with only a lower velocity, taking more time to go near the bottom.     

Context-dependent effects of bluegill in experimental mesocosm communities

Most knowledge of direct and indirect effects of zooplanktivorous fish has come from studies in which a treatment with a zooplanktivore is compared to a fishless control. However, effects of a zooplanktivore may be different in the presence of other fish species because the other fish have direct and indirect effects that may alter the effects of the zooplanktivore in question. To test this hypothesis, we conducted a tank mesocosm experiment of 2×2 factorial design in which the presence and absence of bluegill (Lepomis macrochirus) were cross-classified with the presence and absence of a fish assemblage composed of western mosquitofish (Gambusia affinis) and common carp (Cyprinus carpio). The presence of bluegill decreased Daphnia, Ceriodaphnia, cyclopoid copepodids, calanoid copepodids, copepod nauplii, amphipods, gastropods, and notonectids. Daphnia, Ceriodaphnia, cyclopoid copepodids, copepod nauplii, gastropods, notonectids, Najas, and Chara were decreased and herbivorous rotifers, turbidity, chlorophyll a, total nitrogen and total phosphorus were increased in the presence of the fish assemblage. Significant bluegill×fish assemblage interaction effects were detected for Daphnia, Ceriodaphnia, cyclopoid copepodids, copepod nauplii, gastropods, and notonectids. Analysis of simple effects for these response variables revealed that all significant bluegill effects in the absence of the fish assemblage were not significant in the presence of the fish assemblage. Our results indicate that the effects of bluegill may be context dependent, or dependent upon the presence of other trophically similar fish species. Received: 3 November 1998 / Accepted: 24 September 1999     

Vernal abundance,structure and development of epipelagic copepod populations of the eastern Weddell Sea (Antarctica)

Summary Pelagic copepod populations under the pack ice of the Antarctic Weddell Sea were sampled with a 50 m net between October 2 and December 7, 1986, to study their abundance and developmental stage composition before and at the onset of the vernal phytoplankton bloom. Subadult stages and adult females were incubated to estimate rates of development and egg production. Copepod densities in the upper 200 m were highest for the small-sized species Oithona similis, Oncaea curvata and Ctenocalanus citer. About 95% of the copedite stages belonged to these species, dominated by Oithona. The copepodids were outnumbered by the nauplii in all species, except in Oncaea. The stage distribution in the small-sized species was bimodal with peaks in N3 and C4. The larger species Calanus propinquus and Metridia gerlachei showed peaks in nauplii only. Eggs were relatively abundant in all small and large species. Animals smaller than 1 mm not only were more abundant than the larger ones, but also had a higher total potential respiration. Eggs were produced by incubated females in sea water virtually without food at 0°C. Eggs hatched, and Oithona nauplii developed at a rate of about 7 days per stage. Copepodite stages did not develop significantly. Reproduction in the most abundant species commonly occurred before the algal spring increase when food levels were very low. Maintenance of a stable stage distribution at the expense of a high juvenile mortality seemed to be characteristic for the overwintering strategy of Antarctic copepods.     

The biology and morphology of the marine harpacticoid copepod Heteropsyllus nunni Coull, during encystment diapause

Heteropsyllus nunni Coull, a meiobenthic harpacticoid copepod is the marine crustacean to undergo a state of diapause within a cyst. A 12 month field study indicated H. nunni adults reached peak population densities in winter, with nauplii maturing in the spring, becoming adults by April or May.At the last stage of development, a mature but unmated adult, they begin to prepare for encystment diapause. The copepods remain within their cyst in a state of diapause for 3–4 months during the summer only. Studies on the effects of temperature and photoperiod suggested that these two environmental cues are not crucial for induction or termination of diapause. Low temperature delayed development and time to encystment, while high temperatures accelerated development, making the time to encystment shorter. There were males than females in the cysts in laboratory experiments. Upon excystment, the copepods mate, and females begin egg production within one week. Adults that have excysted and mated die after a few weeks of active reproductive effort. Nauplii go on to mature and begin the univoltine diapause/reproductive cycle.The copepods prepare for dormancy in two ways: they begin to produce and store two types of secretory products to be used in cyst construction; then they produce large quantities of lipid to be used as a nutrient supply throughout diapause. Histochemistry of the cyst-building material indicated the lower urosome is full of two chemically different products. Dorsally, there is a storage sac of proteinaceous material. The ventral sac of secretory product is a mucopolysaccharide. The copepod builds the spherical cysts in a matrix of small and large sand grains. The cysts fit tightly around the ventral portion of the animal in the its flexed position: however, there is a large space between the cyst and the sides of the copepod.Biochemical analysis of the cyst showed it is composed of an amino acid complex similar to collagenous material. Scanning electron microscopy revealed a complex of large cuticular pores located in the lower urosome and caudal rami. There are specific pores for secretion of the two cyst-building products.     

Experimental studies on the development of Lernaeocera branchialis (Copepoda: Pennellidae): population processes from egg production to maturation on the flatfish host

Quantitative population dynamical information derived from laboratory- and field-based experiments is provided for the fish-parasitic copepod, Lernaeocera branchialis, infecting flounder (Platichthys flesus) and whiting (Merlangius merlangus). Adult, post-metamorphosis females from whiting can produce more than one set of egg-strings. The mean number of eggs in each egg-string pair was 1445. At 10 °C these eggs took about 12.7 d after extrusion before hatching of NI nauplii began. Hatching took up to 12 days to be completed with an exponentially declining pattern of output over this period. In the laboratory about 44% of the egg-string egg population successfully passed through the NI to NII nauplius moult and the NII to copepodid moult to produce infective copepodids, a process lasting about 2 d. The non-feeding copepodids had a maximum survival time at 10 °C of 18 d, with a time to 50% survival of 7.5 d. In laboratory infection experiments at 10 °C, copepodids infected flounder and passed through all their developmental stages to adulthood and copulation in a minimum of 25 d. Field experiments on the seabed off Lowestoft in June 1987 with a sea temperature of about 16 °C suggested that the developmental period in those conditions could be as short as 11 d. Previously uninfected flounder in the field experiments became naturally infected with copepodids at a mean rate of not less than 30 parasites per fish d^–1.     

Feeding rates and selectivity among nauplii, copepodites and adult females of Calanus finmarchicus and Calanus helgolandicus

This study focuses on selective feeding by developmental stages of two oceanic copepods, Calanus finmarchicus and Calanus helgolandicus from nauplii to adults. A mixture of four algal species of different biochemical composition, Prorocentrum nanum (dinoflagellate), Thalassiosira minima (diatom), Rhodomonas baltica (cryptophyte) and Dunaliella tertiolecta (chlorophyte), added in an equal biovolume, was used in three different experimental set-ups. In set-up 1 the algal species were present as single cells of similar size (14 μm). In set-up 2 the diatom T. minima was present in chains of two or three cells and was therefore larger than the other algae, while the biovolume of all species remained the same. In set-up 3, the diatom T. minima was excluded from the mixture. Feeding selectivity of the copepods was assessed in relation to the quality of the algal species expressed in terms of carbon and nitrogen content, fatty acid composition, and chain length of the diatom. The results show that younger stages and adult females of C. finmarchicus and C. helgolandicus did not show a preference for an algal species when the algae were of similar size. In the feeding experiments where the diatoms were offered as chains, both copepod species showed a selective behaviour only on the basis of algal size. Individual ingestion rates increased from 0.4 to 0.7 μg C day^–1 for nauplii of both species to 5 μg C day^–1 for adult females of C. helgolandicus to 12 μg C day^–1 for C. finmarchicus. Individual filtration rates ranged from 5 ml day^–1 for C. finmarchicus nauplii to 70–98 ml day^–1 for adult females, and from 3 ml day^–1 for C. helgolandicus nauplii to 35–46 ml day^–1 for adult females. Ingestion and filtration rates per unit body carbon decreased gradually in both copepod species with increasing body carbon. The daily ingested amount of food decreased for C. finmarchicus from 124–134% of the body carbon for nauplii to 19% of the body carbon for adult females, and for C. helgolandicus from 117–137% of the body carbon for nauplii to 13–26% of the body carbon of adult females. Electronic Publication     

Ultrastructure and development of the nephridia in Anaitides mucosa (Annelida,Polychaeta)

Different developmental stages (trochophores, nectochaetae, non-mature and mature adults) of Anaitides mucosa were investigated ultrastructurally. A. mucosa has protonephridia throughout its life; during maturity a ciliated funnel is attached to these organs. The protonephridial duct cells are multiciliated, while the terminal cells are monociliated. The single cilium is surrounded by 14 microvilli which extend into the duct lumen without coming into any contact with the duct cells. Corresponding ultrastructure and development indicate that larval and adult protonephridia are identical in A. mucosa. Differences between various developmental stages can be observed only in the number of cells per protonephridium. A comparison between the funnel cells, the cells of the coelothel and the duct cells reveals that the ciliated funnel is a derivative of the duct. Due to the identical nature of the larval and postlarval protonephridia, such a funnel cannot be a secondary structure. In comparison with the mesodermally derived metanephridial funnel in phoronids it seems likely that the metanephridia of annelids and phoronids evolved convergently.     

Life cycle of Boeckella poppei Mrazek and Branchinecta gaini Daday (King George Island, South Shetlands)

In Lake Wujka, a shallow, polymictic Antarctic lake situated at 15 m from the seashore, several yearly cohorts occur of the copepod Boeckella poppei and one of the fairy shrimp Branchinecta gaini. There is circumstantial evidence that the two species compete for food, and perhaps adult fairy shrimp feed on the nauplii of the copepod. Both species are positively influenced by a measure of salinity. However, when autumn storms massively sweep seawater into the lake, all fairy shrimp are wiped out; no hatching occurs until next spring. In B. poppei, some nauplii and copepodites survive or hatch after the salt flows out of the lake. This is an advantage to the copepod that may balance its coexistence with the shrimp. Its cycle is, however, aborted by the freezing of the lake. In contrast to many other Antarctic lakes, the life cycle of the crustaceans is therefore controlled by salinity rather than freezing.     

Toxicity of the dinoflagellate Gambierdiscus sp. toward the marine copepod Tigriopus japonicus

The toxic dinoflagellate Gambierdiscus sp. has recently been observed in temperate areas in the Southern Sea of the Republic of Korea. The purpose of this study was to examine the toxicity of Gambierdiscus sp. toward the marine copepod Tigriopus japonicus. T. japonicus adult females and nauplii were exposed to various concentrations of algae or culture medium to assess toxicity and analyze gene expression patterns in the copepod. Based on the toxicity tests, Gambierdiscus sp. increased the mortality of nauplii and the immobility of adult females. The survival and mobility of T. japonicus were not affected by culture medium lacking Gambierdiscus sp. cells in the toxicity test. However, based on the analysis of gene expression in the copepod, exposure of the copepod to culture medium affected the expression of stress or detoxification-related genes. Further studies to identify toxins in Gambierdiscus sp. are required to increase our understanding of dinoflagellate toxicity.     

Changes in carotenoid composition in different developmental stages of copepods: Pseudocalanus acuspes Giesbrecht and Acartia spp.

Carotenoid contents and composition were studied in three successivedevelopmental groups of copepod populations from the southernBaltic Sea. Samples were collected with >80% prevalence ofany one of three developmental groups: (i) nauplii, (ii) copepodidsI–III and (iii) copepodids IV–V and also adultsof Pseudocalanus acuspes Giesbrecht and Acartia spp. (Copepoda:Calanoida). Samples were collected over the years 1991–1999from May to September at six sampling sites in the southernBaltic Sea. The carotenoids were analyzed by means of reverse-phasehigh-performance liquid chromatography (RP-HPLC). The mean totalcarotenoid concentrations (astaxanthin, astaxanthin esters,canthaxanthin and other carotenoid unidentified and derivedfrom the diet) found in three groups of developmental stagesfor P. acuspes were 556 µg g^-1 dry weight (d.w.) in naupliarstages, 791 µg g^-1 in copepodids I–III, and 868µg g^-1 in copepodids IV–V and adults. The averagecarotenoid concentrations in Acartia spp. were 619, 764 and872 µg g^-1 d.w., respectively. Significant changes wereobserved in the proportion of carotenoids of separate developmentalgroups. Astaxanthin and canthaxanthin occurred in all developmentalgroups, with the evident dominance of astaxanthin. However,copepodid groups I–III, and especially IV–V includingadults, showed an increased proportion of astaxanthin esters.This suggests that astaxanthin is the main active carotenoidin copepod metabolism. It acts most probably as an efficientfree radical quencher and may be involved in rapid metabolismof stored lipids in mostly unfeeding young nauplii. The feedingstages (late nauplii and copepodids I–III and IV–V)are evidently able to metabolize this pigment by esterificationand further degradation.     

Haplostomides hawaiiensis,new species (Copepods: Cyclopoida: Ascidicolidae), associated with the ascidian Polyclinum constellatum at Honolulu,Hawaii

An ascidicolid copepod, Haplostomides hawaiiensis, is described on the basis of females obtained from the compound ascidian Polyclinum constellatum collected in Keehi Lagoon, Honolulu, Hawaii. The features that separate H. hawaiiensis from other species of the genus include an elongate body with dorsal curvature and indistinct tagmosis, and a combination of characters of cephalic appendages (antenna with 2 spines, mandible with 3 setae, maxillule with 7 setae and maxilla with 2 setae) and thoracic appendages (legs 1–4 with 1 seta on protopod and 2 spines and 1 seta on exopod). Haplostomides hawaiiensis appears to be most closely related to H. hibernicus (T. & A. Scott, 1895) which occurs in compound ascidians from European seas. This is the first copepod associate of an ascidian to be reported from Hawaii.     

Motion behavior of nauplii and early copepodid stages of marine planktonic copepods

The goal of this study was to quantify periods of activity andvelocities of late naupliar and early copepodid stages of planktoniccopepods occurring regularly on the southeastern continentalshelf of the USA. We obtained quantitative information on eightspecies, including adult females of Oithona plumifera. All studieswere conducted at food concentrations near or above satiationlevels. Activities ranged from 0.85% (adult females of O.plumifera)to 100% of time (nauplii and copepodids of various calanoidspecies). Motion velocities (excluding escape motion) coveredmore than one order of magnitude: from 0.39 mm s^–1 fornauplii of Temora stylifera to 5.24 mm s^–1 for naupliiof Oncaea mediterranea. Ranges of activities of species rangefrom occasional for early juveniles to adult females of O.plumiferato 100% for the same range of T.stylifera, the latter creatinga feeding current from N III onwards, the former not at all.Of notable interest is Centropages velificatus which moves intermittentlyas a late nauplius, continuously as an early copepodid and intermittentlyas an adult. All observed calanoid late nauplii and copepodidsmove in three dimensions, excluding copepodids of the shelfbreak/oceanicParacalanus aculeatus. The results indicate not only significantdifferences in motion behavior between cyclopoids and calanoids,but also between calanoid species. Yet, some calanoid speciesshow little ontogenetic changes at all.     

Effect of food level on the growth and survival of laboratory-reared larvae of bay anchovy (anchoa mitchilli Valenciennes) and scaled sardine (harengula pensacolae Goode & Bean)

The effect of food levels on the growth and survival of laboratory-reared larvae of scaled sardines (Harengula pensacolae Goode & Bean) and bay anchovies (Anchoa mitchilli Valenciennes) was determined. Wild zooplankton, most of which was copepod nauplii and copepodids, was used as food. H. pensacolae larvae grew 0.80 mm/day at a low (444 organisms/1) and high food level (1324 organisms/1). Survival was appreciably better (14.5%) at the low than (3.5%) at the high food level, 23 days after hatching. A. mitcchilli larvae grew faster at medium and high food concentrations. Average growth rates for three replicated experiments were 0.48 mm/day at low food (621–692 organisms/1), 0.50 mm/day a t medium food (1330–1688 organisms/1), and 0.54 mm/day at high food (2811–3323 organisms/1). Survival of A. mitchilli larvae, 20 to 28 days after hatching, ranged from 0.0 to 17.8% and was better at medium and high food levels than at the low food level. The resultsng indicate that 500 to 1000 copepod nauplii and copepodids/1 provided adequate food for rearing H. pensacolae but 1500–2000/1 were consistently required to rear A. mitchilli.     

Diel vertical migration by males,females, copepodids and nauplii in a limnetic population of diaptomus (Copepoda)

Diel vertical migration (DVM) by the sexes and life history stages of the copepod Diaptomus novamexicanus (Herrick) was investigated during two summers in a subalpine lake (Castle Lake, California, USA). Migratory behavior was quantified repeatedly by 1300 and 21–2200 hr sampling in the epilimnion. Population density and vertical distribution were estimated routinely at 1300 hr and at 2200 hr on selected dates. Males had greater night to day biomass differentials and ratios than females. The sexes showed comparable DVM participation (% of total biomass migrating) and amplitudes of migration (distance of ascent). Copepodids displayed higher participation, night/day ratios and amplitudes than younger copepods, but adult participation differed only from that of nauplii. Active participation in high amplitude ascents thus increased with age and was greatest in adult males. Analysis suggests that such relative behaviors probably occur in most limnetic Diaptomus, and that the magnitude of ascent into the epilimnion is associated with daytime vertical distribution.     

New species of Ergasilus von Nordmann, 1832 (Copepoda: Ergasilidae) from the gills of freshwater fishes in Patagonia,Argentina

Systematic Parasitology - A new parasitic copepod, Ergasilus yandemontei n. sp., is described based on 10 adult females collected from the gills of the atherinid silverside Odontesthes hatcheri...     

Life history traits of the tropical freshwater copepod Mesocyclops longisetus (Crustacea: Copepoda)

The life history traits of a tropical copepod population of Mesocyclops longisetus were measured in the laboratory under defined and controlled culture conditions. Egg duration times at 25 and 30 °C were 42.1 and 29.6 hours, respectively. Development times from nauplius I to adult females and males were 22.3 and 18.3 days, respectively. Body sizes (cephalothorax length) of adult females and males were 1.00 and 0.60 mm, respectively. Clutch size averaged 70.2 eggs. Comparison of these traits with those of its congeners reveals considerable life history variation in the genus Mesocyclops. Lipid analyses of early stage eggs showed that the percent composition of triglycerides to total lipid in M. longisetus (22%) is similar to that of Mesocyclops edax (17%) and daphniid cladocerans, but only half that of a marine calanoid copepod. We suggest that zooplankton may adapt to different resource environments by either adjusting their egg size or altering the proportion of lipid that is readily available for metabolic energy of embryos and nauplii. The diverse arrays of zooplankton life histories may be explained in part by adaptations in lipid metabolism in response to the level and predictability of resources.     

Prey capture capabilities by juveniles of the false percula clownfish (Amphiprion ocellaris) fed calanoid nauplii vs. adults

The reaction field of juvenile false percula clownfish (Amphiprion ocellaris) were studied when fed two different developmental stages (nauplii and adults) of the calanoid copepod Pseudodiaptomus annandalei. As the copepods undergo ontogenetic development, their morphologic and kinematic traits changes and may therefore influence the ability of the clownfish to identify and capture prey. The fish reacted to nauplii at shorter distances (15.2 ± 7.1 mm) compared to adult copepods (28.0 ± 9.8 mm). However, the fish reaction angle was significantly wider when offered nauplii (?25.3° to 64.4°) compared to adult copepods (?14.2° to 36.7°). This resulted in an equivalent attack rate on both prey items. Hence, even though nauplii are less apparent at greater distances, the fish counteract this partly by reacting to smaller prey items at a wider range. However, the carbon-specific ingestion rate was higher when offered adult copepods, which suggests adult copepods are a more rewarding prey.     

Characteristics of the reproductive cycles and development times of Copidodiaptomus numidicus (Copepoda: Calanoida) and Acanthocyclops robustus (Copepoda: Cyclopoida)

The female reproductive cycles of the calanoid copepod Copidodiaptomus numidicus and the cyclopoid Acanthocyclops robustus were characterized by periodic changes in the oviducts. Females oscillated between a gravid and a non-gravid condition, and the whole cycle was strongly dependent on temperature. In both species, the maturation of new oocytes took place when the old egg sacs were still being carried, thus ensuring a rapid clutch succession. The embryonic and post-embryonic development duration of C.numidicus and A.robustus was investigated at five and four different temperatures, respectively. Embryonic development time was related to oviductal cycle duration at each temperature to estimate the minimum amount of non-ovigerous females expected under non-limiting mate and food conditions. Copidodiaptomus numidicus females spent 35-19% of the cycle without carrying eggs at 7-20C. Acanthocyclops robustus females spent 31-33% of the cycle without carrying eggs at the same temperature interval. Both naupliar and copepodid durations were inverse functions of temperature with nauplii developing at a faster rate than copepodids. In general, A.robustus showed reproductive advantage over C.numidicus due to a faster oviductal cycle and shorter embryonic and post-embryonic development times. However, the influence of food quality may be crucial. Acanthocyclops robustus copepodid development may lengthen well beyond the corresponding development stages of C.numidicus when growing only on algae. This retardation of development affects especially the later development stages of the cyclopoid.      

Inhibition of copepod feeding by exudates and transparent exopolymer particles (TEP) derived from a Phaeocystis globosa dominated phytoplankton community

We investigated if (1) dissolved compounds excreted by Phaeocystis globosa and (2) transparent exopolymer particles (TEP) formed from carbohydrates excreted into the water affect the feeding of nauplii and females of the calanoid copepod Temora longicornis during a P. globosa bloom. Copepod grazing on the diatom Thalassiosira weissflogii in the presence of these possible grazing deterrents was measured during three successive weeks of a mesocosm study, simulating the development of a P. globosa bloom. Our results demonstrate no indication for the presence of feeding deterrents in the dissolved phase, but a strong inhibitory effect of transparent exopolymer particles (TEP) on the consumption of algae by both nauplii and adult copepods. The inhibitory effect of TEP was connected to the accumulation of DOM during the progress of the bloom. We suggest that a reduction in the grazing pressure of zooplankton may increase the survival of the liberated single cells during disruption of colonies and allow seeding populations to persist. Furthermore, P. globosa reduces the trophic efficiency of the food web not only by withdrawal of its colonies from grazing but also by a relaxation of the grazing pressure on co-occurring phytoplankton and by alteration of the food web structure via TEP production.