Omnivorousness in marine planktonic copepods

Adult females of Centropages furcatus and Temora stylifera hadsignificantly different feeding behaviours. At equal averagefood concentrations, C. furcatus ingested more nauplii of thecalanoid copepod Pseudodiaptomus coronatus and fewer diatomsthan T. stylifera. At 20°C and 2.0 µg of nitrogen.1.1.^–1of each food source, C. furcatus ingested 17.2% and 4.4% ofits body nitrogen daily as nauplii and diatoms, respectively.Ingestion rates of the same food sources by T. stylifera were5.1% and 14.5%, respectively. At the environmental concentrationssimulated in these experiments, the presence of diatoms didnot affect the ingestion of nauplii nor did the presence ofnauplii affect diatom ingestion. The behaviour of predator andanimal prey largely dictates the types and quantities of preyingested.     

Evolution of bioluminescence in marine planktonic copepods

Copepods are the dominant taxa in zooplankton communities of the ocean worldwide. Although bioluminescence of certain copepods has been known for more than a 100 years, there is very limited information about the structure and evolutionary history of copepod luciferase genes. Here, we report the cDNA sequences of 11 copepod luciferases isolated from the superfamily Augaptiloidea in the order Calanoida. Highly conserved amino acid residues in two similar repeat sequences were confirmed by the multiple alignment of all known copepod luciferases. Copepod luciferases were classified into two groups of Metridinidae and Heterorhabdidae/Lucicutiidae families based on phylogenetic analyses, with confirmation of the interrelationships within the Calanoida using 18S ribosomal DNA sequences. The large diversity in the specific activity of planktonic homogenates and copepod luciferases that we were able to express in mammalian cultured cells illustrates the importance of bioluminescence as a protective function against predators. We also discuss the relationship between the evolution of copepod bioluminescence and the aspects of their ecological characteristics, such as swimming activity and vertical habitat.     

Ciliates as a food source for marine planktonic copepods

Copepods of the genusEurytemora, isolated from the Patuxent River, a tributary of Chesapeake Bay, were fed suspensions of the ciliateUronema isolated from the Rhode River, a subestuary of Chesapeake Bay. Grazing by copepods was determined by the decrease in numbers of ciliates, which were monitored by both direct counting and particle size analysis. Results from both methods of analysis showed significant reduction in the numbers ofUronema in the suspension whenEurytemora was present. Survival of copepods with ciliates added as food source was significantly longer than without ciliates. Analysis of field samples collected in the fall showed that ciliates comprised approximately 20% of the total plankton biomass at selected sampling sites. The results of the laboratory and field studies indicate that copepods can feed on ciliates and suggest that, in nature, ciliates may comprise an important source of food for copepods.     

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.     

Egg-brooding, body size and predation risk in planktonic marine copepods

We investigated the interacting effects of copepod body size and the presence or absence of egg masses on the risk of predation by a visual predator. We conducted selection experiments involving three-spined sticklebacks (Gasterosteus aculeatus) and copepods ranging in body mass from 0.5 to 740 μg C: Oithona similis, Corycaeus anglicus, Pseudocalanus newmani, P. moultoni, Pseudodiaptomus marinus, and Paraeuchaeta elongata. We found that sticklebacks selected ovigerous females of the two smallest-bodied species of copepods (Oithona similis and Corycaeus anglicus). In contrast, the fish showed no significant selection for ovigerous females of the remaining, larger-bodied species. Unexpectedly, egg mass position (i.e., in a ventral, dorsal or lateral location on the urosome) appeared to influence predation risk more than did body size, resulting in higher predation risk for the cyclopoid and poecilostomatoid species than for the calanoid species we tested. Although the sticklebacks showed no statistically significant preference for ovigerous females of any of the four calanoid species, for each species the overall proportion of ovigerous females ingested was slightly greater than 0.50. Thus, whether body size influences the susceptibility of egg-brooding calanoid copepods to predation remains an open question. Received: 24 August 1998 / Accepted: 6 July 1999     

Histomorphological changes in the reproductive condition of parasitized marine planktonic copepods

Changes in the internal morphology of parasitized copepods sampledin the Gulf of Naples from 1986 to 1988 were investigated usingboth light and transmission electron microscopy. The most commonand devastating form of infection was due to the parasitic dinoflagellateSyndinium, which induced gross modifications in the internaland external morphology of the host population. The parasitereduced fecundity and longevity of infected individuals, whichmainly included juvenile and female populations of the copepodParacalanus parvus. Infection rates for this species were ashigh as 12% for. juveniles and 13% for adult females for theperiod investigated. Infection by the parasitic dinoflagellateBlastodinium induced less negative effects on the reproductivebiology of their hosts. This pathogen did not lead to sexualcastration and oogonal development appeared normal. Other formsof infestation were rare and included infections due to protozoa,fungi and bacteria. ^*Died tragically at sea during a cruise on December 15, 1988     

A metagenetic approach for revealing community structure of marine planktonic copepods

Marine planktonic copepods are an ecologically important group with high species richness and abundance. Here, we propose a new metagenetic approach for revealing the community structure of marine planktonic copepods using 454 pyrosequencing of nuclear large subunit ribosomal DNA. We determined an appropriate similarity threshold for clustering pyrosequencing data into molecular operational taxonomic units (MOTUs) using an artificial community containing 33 morphologically identified species. The 99% similarity threshold had high species‐level resolution for MOTU clustering but overestimated species richness. The artificial community was appropriately clustered into MOTUs at 97% similarity, with little inflation in MOTU numbers and with relatively high species‐level resolution. The number of sequence reads of each MOTU was correlated with dry weight of that taxon, suggesting that sequence reads could be used as a proxy for biomass. Next, we applied the method to field‐collected samples, and the results corresponded reasonably well with morphological analysis of these communities. Numbers of MOTUs were well correlated with species richness at 97% similarity, and large numbers of sequence reads were generally observed in MOTUs derived from species with large biomass. Further, MOTUs were successfully classified into taxonomic groups at the family level at 97% similarity; similar patterns of species richness and biomass were revealed within families with metagenetic and morphological analyses. At the 99% similarity threshold, MOTUs with high proportions of sequence reads were identified as biomass‐dominant species in each field‐collected sample. The metagenetic approach reported here can be an effective tool for rapid and comprehensive assessment of copepod community structure.     

Elemental and isotopic fractionation of carbon and nitrogen by marine, planktonic copepods and implications to the marine nitrogen cycle

Particle-grazing copepods, primarily Temora longicornis andT. stylifera, and seawater with natural particles were collectedfrom the northwest Gulf of Mexico. Control and ammonium-enrichedaliquots of seawater were incubated in triplicate for 2 days,copepods added and the incubation continued for 2 days. Analyseswere made of dissolved nutrients (nitrate, ammonium and phosphate),suspended particles (chlorophyll a and phaeopigments, C, N,     

Propagation of planktonic copepods: production and mortality of eggs

Data on fecundity and egg mortality of neritic copepods were collected in various seasons, areas and under various hydrographical conditions. On a seasonal basis variations in fecundity (F) were related to temperature rather than to the abundance of phytoplankton (P). However, a strong correlation between F and P was evident when water column stability varied horisontally or temporally (i.e. at a tidal front or subsequent to a storm). Estimated specific egg-mortalities were variable and occasionally very severe, up to 9.1 d^–1, implying that down to 10^–4% of the eggs survive to hatching. The implications for phenology and distribution of copepods are discussed.     

Feeding rates of planktonic copepods from a tropical sea

The feeding rates of the marine planktonic copepods, Eucalanus subcrassus Giesbrecht, Tortanus gracilis (Brady), Calanopia elliptica (Dana) (both male and female), Temora turbinata (Dana), and Paracalanus aculeatus Giesbrecht (only female) from tropical inshore waters have been studied. Newly hatched Artemia nauplii (for Eucalanus, Tortanus, and Calanopia), Dunaliella (for Temora), and Skeletonema (for Paracalanus) were used as food.Feeding rates were measured for a single individual through successive incubations once or twice a day until death to determine changes in feeding rate after collection. Copepods survived from a few days to three weeks. In general, feeding rates varied from day to day, but were less variable than the differences between day and night rates. In some cases, feeding rate consistently decreased up to the death of the copepod. Daily ration, estimated in terms of percentage body weight, was in the range of 28–329 ‰ Using the results together with the those of other workers, gives the relation between daily ration (Y, % body weight) and body weight of copepods (X; μg dry weight) at 20 °C as, logY = 2.531?0.377 logX.Copepods given Artemia nauplii as food killed more nauplii than were eaten. This phenomenon, tentatively called ‘over-hunting’, is possibly an important feeding behaviour for carnivorous copepods.     

Planktonic marine copepods and harmful algae

Marine planktonic copepods are important grazers on harmful algae (HA) species of phytoplankton, and copepods are major entry points for vectorial intoxication of pelagic food webs with HA toxins. Previous reviews (Turner and Tester, 1997, Turner et al., 1998a, Turner, 2006) summarized information on HA interactions with zooplankton grazers, and vectorial intoxication of pelagic food webs, up through approximately 2005. Accordingly, this review will address primarily studies published during the last decade. It will concentrate on generic issues in the developing field of HA:grazer interactions, such as the extent to which HA toxins serve as copepod grazing deterrents, induction of HA grazing deterrents by exposure to copepods, copepod selective feeding to avoid ingesting HA taxa versus non-selective feeding on HA taxa, possible biogeographic aspects of the effects of HA toxins on copepods, impact of copepod grazing on HA bloom development and termination, the role of copepods as entry points for vectorial intoxication of pelagic food webs with HA toxins, and possible reasons and remedies for the highly-variable and conflicting results reported for many studies of copepod grazing on various HA species.     

Ecological relationships between Vibrio cholerae and planktonic crustacean copepods.

Strains of Vibrio cholerae, both O1 and non-O1 serovars, were found to attach to the surfaces of live copepods maintained in natural water samples collected from the Chesapeake Bay and Bangladesh environs. The specificity of attachment of V. cholerae to live copepods was confirmed by scanning electron microscopy, which revealed that the oral region and egg sac were the most heavily colonized areas of the copepods. In addition, survival of V. cholerae in water was extended in the presence of live copepods. Attachment of viable V. cholerae cells to copepods killed by exposure to -60 degrees C was not observed. Furthermore, survival of V. cholerae was not as long in the presence of dead copepods as in the live copepod system. A strain of Vibrio parahaemolyticus was also seen to attach to copepod surfaces without effect on survival of the organism in water. The attachment of vibrios to copepods was concluded to be significant since strains of other bacteria, including Pseudomonas sp. and Escherichia coli, did not adhere to live or dead copepods. Attachment of V. cholerae to live copepods is suggested to be an important factor of the ecology of this species in the aquatic environment, as well as in the epidemiology of cholera, for which V. cholerae serovar O1 is the causative agent.     

Taxonomic studies on brackish copepods in Korean waters--II. Ontogeny and phylogeny of appendages in copepodid stages of Tortanus derjugini Smirnov, 1935 (Copepoda, Calanoida)

The development of all copepodid stages of a brackish calanoidcopepod Tortanus (Eutortanus) derjugini from western Korea isinvestigated. Homologies of segmentation and setation in adultcharacters are traced through all copepodid stages. The fourthand fifth pedigers are secondarily fused at the moult to theadult. Antennulary setation shows sexual dimorphism first atstage IV. The exopod of leg 1 is divided into two segments withoutaddition of any element at the moult to stage II, while an innerseta on the first segment is newly added at the moult to stageIII. This pattern differs from the basic pattern proposed forother copepods, in which the division of the exopod is precededby the formation of an element or both occur at the same time.In the fifth leg of the male some elements are suppressed duringthe final moult into the adult. The ontogenetic analyses indicatethat adult characters of some subgenera of Tortanus are expressedby their suppressions at the developmental phases. Suppressionof elements is important for the evolution of Tortanus.     

Late Miocene—early Pliocene planktonic foraminiferal biostratigraphy and paleoceanography of low-latitude marine sequences

Plate reorganization and development of polar glaciation are closely associated with the changing climatic conditions of the Cenozoic. The planktonic foraminiferal fauna in three low-latitude DSDP sites (224, Ninety East Ridge; 317B, Manihiki Plateau; 366A, Sierra Leone Rise) has been examined to determine how these regions responded to the late Miocene climatic cooling that has been previously observed in high-latitude regions. It has been proposed that an expansion of Antarctic glaciation and a resultant eustatic regression are associated with this cooling, with the latter being at least partially responsible for the Messinian “salinity crisis” in the Mediterranean.Only one of the sites, 366A, shows any significant faunal change during the late Miocene and early Pliocene (approximately 8-3 Ma). During the late Miocene, there is a decrease in the abundance of species considered to be tropical-subtropical, suggesting the incursion of a cool water mass into the Sierra Leone Rise region during this time. The lack of any major faunal changes at Sites 214 and 317B indicates that the water masses of the low-latitude regions of the Indo-Pacific were relatively unaffected by this cooling in the late Miocene.The three sites also show no evidence for a change in the level of the CCD between the late Miocene and early Pliocene; however, this is most likely due to their equatorial position and shallow water depth. Higher-latitude sites from both the Atlantic and the Pacific reveal a definite shoaling of the CCD during the late Miocene.     

Neural development in the planktonic and early benthic stages of the palinurid lobster Jasus edwardsii

Jasus edwardsii has eleven phyllosoma larvae that are morphologically adapted for a plank‐tonic existence. These are followed by a puerulus stage which, although it has the general body form of the adult, is also adapted for swimming. The next moult is the first of the truly benthic juvenile stages. Using transmission electron microscopy we compared transverse sections of connectives in the ventral nerve cord of the first phyllosoma, the puerulus and the first post‐puerulus stage at a number of points. We found that the anterior cord of the phyllosoma has approximately the same number of neurons as the puerulus. This has implications for our understanding of crustacean larval sensory systems because previous work shows that most fibres in the adult nerve cord are sensory. Fibre counts in the post‐puerulus increase in the expected manner. The cord structure of the puerulus and post‐puerulus is the same as that found in the adult whereas that of the early phyllosoma is not.     

The escape behavior of marine copepods in response to a quantifiable fluid mechanical disturbance

The threshold shear values needed to elicit the escape reactionto a quantifiable fluid mechanical disturbance were comparedbetween five free-swimming oceanic copepod species. The resultsindicate a significant difference in the threshold for differentspecies of copepods and between different age groups withina single species. In general, animals captured from more energeticregimes required a higher threshold than those captured frommore pacific locations. Labidocera madurae required the highestshear values with 51.5 s^–1 for 50% of the animals testedto elicit an escape reaction (S₅₀). Acartia tonsa and Euchaetarimana, in contrast, were behaviorally the most sensitive requiringan S₅₀ of only 1.5 and 4.1 s^–1, respectively, to initiatean escape reaction. Pleuromamma xiphias and Oithona requiredintermediate shear values with an S₅₀ of 7.2 and 8.1 s^–1.When compared to literature values, the threshold needed toelicit an escape reaction was consistently higher than averageenvironmental shear values. Threshold shear values also variedsignificantly with developmental stage. Naupliar stages of A.tonsarequired greater than six times the S₅₀ value required by adultsof the same species. This suggests that the higher vulnerabilityto predation of naupliar stages of copepods may not only reflectinferior escape strength, but may also result from the higherthreshold needed to elicit an escape reaction. This study supportsthe hypothesis that selective feeding patterns exhibited bypredators of copepods may be the result of the differentialbehavioral sensitivities of different species and developmentalstages of copepods.     

Comparative organ differentiation during early life stages of marine fish

The basic developmental mechanisms of teleosts are similar, but there are differences with respect to the timing of developmental events. These events are controlled by genetic and environmental factors. Direct comparisons of organogenesis are complicated due to large variations in egg sizes and incubation temperatures between species. But in general, cultivated small marine pelagic fish larvae originating from rather small eggs (like gadoids, flatfishes, sparids) hatch with a relatively large yolk sac, a larval finfold and subdermal space and under-developed organs. Developmental status at hatch differs between species and the duration of the yolk sac period varies. Main organs and organ systems become functional by first feeding and differentiate during the larval stage and metamorphosis. Species developing directly via large yolk-rich eggs and a long incubation period have a juvenile like morphology and organ functionality at first feeding, sometimes immediately after hatch (like wolffishes). Histomorphological and cell- or organ functional studies of developing embryos and larvae of cultivated species constitute basic information for understanding species-specific events, of utmost importance for improving production protocols. Information is still lacking on early functionality of endocrine and immunocompetent tissues and organs, areas that deserve future focus.     

Feeding, excretion and egg production by individuals and populations of the marine, planktonic copepods, Acartia spp. and Centropages furcatus

Diel variations in vertical distribution, gut pigment content,ammonium excretion and egg production were investigated foradult females of Acartia erythraea and A.pacifica in the verticallymixed Inland Sea of Japan and Centropages furcatus in the stratified,neritic Gulf of Mexico. Gut pigment content and egg productionrate were maximal at night and ammonium excretion was maximalduring the daytime. Neither A.erythraea nor A.pacifica adultfemales showed an apparent diel migration, but the former werehighly concentrated in the surface layer during the afternoon.In contrast, C.furcatus adult females showed a clear diel migration,residing immediately above the bottom during the daytime andbeing concentrated between 10 and 25 m depth during the nighttime.Individual-based data on gut content and excretion and egg productionrates were combined with vertical-distribution data to calculatepopulation values. In the Inland Sea of Japan, the resultantpattern for Acartia spp. reflected the diel variation in physiologicalrates and even distribution of adult females, except for theafternoon, surface aggregation of A.erythraea. In the Gulf ofMexico, the pattern for C.furcatus reflected largely the dielvariation in each rate process and the heterogeneous distributionof adult females in the water column. Elevated nocturnal feedingactivity of these copepods may be due to an endogenous rhythm.The daytime maximum in ammonium excretion and night-time maximumin egg production rate indicated approximate half-day and daytime lags, respectively, after the intake of food until itsconversion into dissolved excreta and released eggs.     

Life in transition: balancing inertial and viscous forces by planktonic copepods

Copepods (1-10 mm aquatic crustaceans moving at 1-1000 mm s(-1)) live at Reynolds numbers that vary over 5 orders of magnitude, from 10(-2) to 10(3). Hence, they live at the interface between laminar and turbulent regimes and are subject to the physical constraints imposed by both viscous and inertial realms. At large scales, the inertially driven system enforces the dominance of physically derived fluid motion; plankton, advected by currents, adjust their life histories to the changing oceanic environment. At Kolmogorov scales, a careful interplay of evenly matched forces of biology and physics occurs. Copepods conform or deform the local physical environment for their survival, using morphological and behavioral adaptations to shift the balance in their favor. Examples of these balances and transitions are observed when copepods engage in their various survival tasks of feeding, predator avoidance, mating, and signaling. Quantitative analyses of their behavior give measures of such physical properties of their fluid medium as energy dissipation rates, molecular diffusion rates, eddy size, and eddy packaging. Understanding the micromechanics of small-scale biological-physical-chemical interactions gives insight into factors influencing large-scale dynamics of copepod distribution, patchiness, and encounter probabilities in the sea.