Electrophoretic survey of genetic variation in Pecten maximus,Chlamys opercularis,C. varia and C. distorta from the Irish Sea

Food selection by young larvae of the gulf menhaden (Brevoortia patronus) was studied in the laboratory at Beaufort, North Carolina (USA) in 1982 and 1983; this species is especially interesting, since the larvae began feeding on phytoplankton as well as microzooplankton. When dinoflagellates (Prorocentrum micans), tintinnids (Favella sp.), and N1 nauplii of a copepod (Acartia tonsa) were presented to laboratory-reared, larval menhaden (3.9 to 4.2 mm notochord length), the fish larvae ate dinoflagellates and tintinnids, but not copepod nauplii. Larvae showed significant (P<0.001) selection for the tintinnids. Given the same mixture of food items, larger larvae (6.4 mm notochord length) ate copepod nauplii as well as the other food organisms. These feeding responses are consistent with larval feeding in the northern Gulf of Mexico, where gulf menhaden larvae between 3 and 5 mm in notochord length frequently ate large numbers of dinoflagellates (mostly P. micans and P. compressum) and tintinnids (mostly Favella sp.), but did not eat copepod nauplii. As larvae grew, copepod nauplii and other food organisms became important, while dinoflagellates and tintinnids became relatively less important in the diet. Since the tintinnids and nauplii used in the laboratory feeding experiments were similar in size as well as carbon and nitrogen contents, the feeding selectivity and dietary ontogeny that we observed were likely due to a combination of prey capturability and larval fish maturation and learning.Contribution No. 5575 of the Woods Hole Oceanographic Institution     

Filtering rates of the juvenile Atlantic menhaden Brevoortia tyrannus (Pisces: Clupeidae), with consideration of the effects of detritus and swimming speed

From July to September 1982 feeding experiments were conducted with 138-mm fork length Atlantic menhaden Brevoortia tyrannus (Latrobe) (Pisces: Clupeidae) to determine their particle size-specific feeding abilities. Monoculture clearing-rate experiments showed that the minimum size of particles filtered, the minimum size threshold, for 138-mm fish is 7 to 9 m. Filtration efficiency for three species of phytoplankton below the minimum size threshold. Pseudoisochrysis paradoxa, Monochrysis lutheri, and Isochrysis galbana, averaged 1.0% (n=14). Tetraselmis suecica, Prorocentrum minimum, and 2-celled Skeleionema costatum, phytoplankton which are larger than the minimum size threshold and smaller than the 20-m upper limit for nanoplankton, were filtered at efficiencies averaging 21% (n=24). S. costatum chains of 3 to 6 cells, prey particles exceeding the size limits of nanoplankton, were filtered at average efficiencies ranging from 22 to 84%. The mean filtration efficiency for Artemia sp. nauplii (San Francisco Bay Brand) of 36% (n=7) was lower than for smaller phytoplankton prey. The presence of detritus at concentrations usually encountered in nature enhanced filtering efficiency and lowered minimum size thresholds at which phytoplankton were retained. For small food particles, filtering efficiency decreased as swimming speed of the menhaden increased. As menhaden grow, their feeding tepertoire shifts to larger planktonic organisms.Contribution No. 1201 Virginia Institute of Marine Science     

Integration of a fish bioenergetics model into a spatially explicit water quality model: Application to menhaden in Chesapeake Bay

Although fish are usually thought of as victims of water quality degradation, it has been proposed that some planktivorous species may improve water quality through consumption of algae and sequestering of nutrients via growth. Within most numerical water quality models, the highest trophic level modeled explicitly is zooplankton, prohibiting an investigation of the effect a fish species may be having on its environment. Conversely, numerical models of fish consumption do not typically include feedback mechanisms to capture the effects of fish on primary production and nutrient recycling. In the present study, a fish bioenergetics model is incorporated into CE-QUAL-ICM, a spatially explicit eutrophication model. In addition to fish consumption of algae, zooplankton, and detritus, fish biomass accumulation and nutrient recycling to the water column are explicitly accounted for. These developments advance prior modeling efforts of the impact of fish on water quality, many of which are based on integrated estimates over an entire system and which omit the feedback the fish have through nutrient recycling and excretion. To validate the developments, a pilot application was undertaken for Atlantic menhaden (Brevoortia tyrannus) in Chesapeake Bay. The model indicates menhaden may reduce the algal biomass while simultaneously increasing primary productivity.     

Vulnerability of the copepod Acartia tonsa to predation by the scyphomedusa Chrysaora quinquecirrha : effect of prey size and behavior

Although scyphomedusae have received increased attention in recent years as important predators in coastal and estuarine environments, the factors affecting zooplankton prey vulnerability to these jellyfish remain poorly understood. Current models predicting feeding patterns of cruising entangling predators, such as Chrysaora quinquecirrha (Desor, 1948), fail to account for the selection of fast-escaping prey such as copepods. Nevertheless, our analysis of gastric contents of field-collected medusae showed that this scyphomedusa fed selectively on the calanoid copepod Acartia tonsa (Dana, 1846) and preferentially ingested adult over copepodite stages. We measured feeding rates in a planktonkreisel while simultaneously videotaping predator–prey interactions. C. quinquecirrha consumed adult A. tonsa ten times faster than copepodites. Differences in prey behavior, in the form of predator–prey encounter rates or post-encounter escape responses, could not account for the elevated feeding rates on adults. Prey size, however, had a dramatic impact on the vulnerability of copepods. In experiments using heat-killed prey, feeding rates on adults (1.5 times longer than copepodites) were 11 times higher than on copepodites. In comparison, medusae ingested heat-killed prey at only two to three times the rate of live prey. These results suggest that during scyphomedusan–copepod interactions, prey escape ability is important, but ultimately small size is a more effective refuge from predation. Received: 26 September 1997 / Accepted: 20 May 1998     

Optimization of exergy and implications of body sizes of phytoplankton and zooplankton in an aquatic ecosystem model

Size appears to be an important parameter in ecological processes. All physiological processes vary with body size ranging from small microorganisms to higher mammals. In this model, five state variables — phosphorus, detritus, phytoplankton, zooplankton and fish are considered. We study the implications of body sizes of phytoplankton and zooplankton for total system dynamics by optimizing exergy as a goal function for system performance indicator. The rates of different sub-processes of phytoplankton and zooplankton are calculated, by means of allometric relationships of their body sizes. We run the model with different combinations of body sizes of phytoplankton and zooplankton and observe the overall biomass of phytoplankton, zooplankton and fish. The highest exergy values in different combinations of phytoplankton and zooplankton size indicate the maximum biomass of fish with relative proportions of phytoplankton and zooplankton. We also test the effect of phosphorus input conditions corresponding to oligotrophic, mesotrophic, eutrophic system on its dynamics. The average exergy to be maximized over phytoplankton and zooplankton size was computed when the system reached a steady state. Since this state is often a limit cycle, and the exergy copies this behaviour, we averaged the exergy computed for 365 days (duration of 1 year) in the stable period of the run. In mesotrophic condition, maximum fish biomass with relative proportional ratio of phytoplankton, zooplankton is recorded for phytoplankton size class 3.12 (log V μm³ volume) and zooplankton size 4 (log V μm³ volume). In oligotrophic condition the highest average exergy is obtained in between phytoplankton size 1.48 (log V μm³ volume) and zooplankton size 4 (log V μm³ volume), whereas in eutrophic condition the result shows the highest exergy in the combination of phytoplankton size 5.25 (log V μm³ volume) and zooplankton size 4 (log V μm³ volume).     

Effects of water motion and prey behavior on zooplankton capture by two coral reef fishes

Water motion is an important factor affecting planktivory on coral reefs. The feeding behavior of two species of tube-dwelling coral reef fish (Chaenopsidae) was studied in still and turbulent water. One species of blenny, Acanthemblemaria spinosa , lives in holes higher above the reef surface and feeds mainly on calanoid copepods, while a second, A. aspera , lives closer to the reef surface, feeds mainly on harpacticoid copepods, and is exposed to less water motion than the first. In the laboratory, these two blenny species were video recorded attacking a calanoid copepod ( Acartia tonsa, evasive prey) and an anostracan branchiopod (nauplii of Artemia sp., passive prey). Whereas A. spinosa attacked with the same vigor in still and turbulent water, A. aspera modulated its attack with a more deliberate strike under still conditions than turbulent conditions. For both fish species combined, mean capture success when feeding on Artemia sp. was 100% in still water and dropped to 78% in turbulent water. In contrast, when feeding on Acartia tonsa, mean capture success was 21% in still water and rose to 56% in turbulent water. We hypothesize that, although turbulence reduces capture success by adding erratic movement to Artemia sp. (passive prey), it increases capture success of Acartia tonsa (evasive prey) by interfering with the hydrodynamic sensing of the approaching predator. These opposite effects of water motion increase the complexity of the predator-prey relationship as water motion varies spatially and temporally on structurally complex coral reefs. Some observations were consistent with A. aspera living in a lower energy benthic boundary layer as compared with A. spinosa: slower initial approach to prey, attack speeds modulated according to water velocity, and lower proportion of approaches that result in strikes in turbulent water.Communicated by P.W. Sammarco, Chauvin     

Functional responses of juvenile herring and sprat in relation to different prey types

The relationship between particulate-feeding rates and prey concentrations (functional response) of juvenile herring and sprat (5–9 cm total length) was investigated in controlled feeding experiments monitored by an underwater camera system. A special tank system was developed allowing the regulation and quantification of low prey concentrations (1–160 L^?1). Non-evasive Artemia nauplii was used as prey to estimate the maximum biting rate of both predators. In contrast, Acartia tonsa with a high escape ability was used as a realistic prey type. Herring and sprat showed a type II functional response for both prey types. Nonlinear mixed effects model revealed no significant difference between the functional responses of both predators, except that herring showed significantly higher biting rates than sprat at A. tonsa concentrations below ~40 L^?1. For both predators feeding rates were significantly higher with Artemia nauplii than with A. tonsa. Video analysis indicated that sprat, unlike herring, is an obligate particulate-feeder.     

Escape responses of copepod nauplii in the flow field of the blue mussel,<Emphasis Type="Italic"> Mytilus edulis</Emphasis>

Bivalves are important grazers on phytoplankton in shallow waters. However, very little is known about their ability to capture actively moving zooplankton. We investigated the escape response and success of early and late nauplii of three copepod species (Acartia tonsa, Temora longicornis and Eurytemora affinis) in the flow field of a blue mussel, Mytilus edulis, using both video observations and incubation experiments. An empirical model was created to describe the spatial distribution of the fluid deformation rate. Nauplii responded with escape jumps at mean fluid deformation rates of 0.6-1.9 s^у. Escape success differed between taxa. T. longicornis was the poorest escaper, while A. tonsa and E. affinis were more efficient and similar to one another. Deformation rates differed in different parts of the flow field, which resulted in differences in escape success between the sectors. Nauplii were caught most often in the sector furthest away from the exhalent siphon, where the deformation rate was the weakest. There the nauplii were unable to detect an escape signal in time to react and flee.     

Quantitative feeding ecology of the hydromedusan Nemopsis bachei in Chesapeake Bay

We determined feeding rates of the hydromedusan Nemopsis bachei L. Agassiz in the mesohaline region of Chesapeake Bay, USA during the spring of 1989 and 1990 from gut contents, digestion rates and abundances of medusae and zooplankton. The medusae consumed primarily copepodites of Acartia tonsa, selecting against naupliar stages. The peak abundance of N. bachei medusae was in April to May, when densities averaged more than 10 m^-3. Medusa densities were similar in both years, but were greatest (maximum of 132 medusae m^-3) along a southern transect sampled only in 1990. At peak densities, N. bachei medusae consumed 30% d^-1 of the copepodite standing stocks, but they consumed <1% d^-1 at the lower densities typical of late May or early June. The predation effects were generally greater than those reported for other hydromedusan species. But even at peak predation, N. bachei medusae could not have controlled or reduced A. tonsa copepod populations, which had a production rate of 85% d^-1 at that time. Medusa feeding rates were highest at nighttime, and were correlated with prey density in the field, but not in the laboratory.Communicated by J. Grassle, New Brunswick     

Critical thermal maxima and upper lethal temperatures for the calanoid copepods Acartia tonsa and A. clausi

Determination of the critical thermal maxima and upper lethal temperatures for Acartia tonsa Dana and A. clausi Giesbrecht revealed the relative temperature tolerance of the two species. A. tonsa exhibited a wider range of acclimation temperatures compared to A. clausi. The response of A. tonsa is related to its ubiquity in Atlantic Ocean estuaries (temperate and tropical), and its capacity to live in temperate waters throughout most of the year. Differences between upper lethal temperatures and critical thermal maxima were very narrow for A. tonsa from tropical and subtropical areas. This suggests that, in warm climates, this copepod lives close to its upper lethal temperature limits.Based in part on a dissertation submitted in partial fulfillment for the Ph.D. degree at the University of Rhode Island.     

Food selection by copepods: discrimination on the basis of food quality

The copepod Acartia tonsa displayed nearly two-fold higher ingestion rates on faster-growing cells of the diatom Thalassiosira weissflogii compared to ingestion rates on slower-growing cells of that species at the same cell concentration. Ingestion rates on slow-growing cells were also enhanced by the addition of cell-free aliquots of algal exudate to the experimental feeding chambers. In addition, the faster-growing algal cells were selectively ingested by the copepod when the two cell types were mixed together in different proportions, indicating that physiological differences between growing cells are a critical factor in the food detection/selection process of zooplankton. Consideration of cell carbon, nitrogen, and protein composition suggests that the copepods are maximizing nitrogenous ingestion (total protein and/or nitrogen). Selectivity for cells with higher protein content results in a higher daily protein ration, even if the selection process results in a decreased rate of ingestion in mixtures of cell types.     

Grazing in juvenile stages of some estuarine calanoid copepods

The grazing of juvenile Eurytemora affinis, Acartia tonsa and A. clausi from the Chesapeake Bay (USA) was investigated using natural particle distributions and freshly caught copepods, live-sorted into stages. Data were analyzed in 110 size channels using an electronic particle counter, and filtering rates (FR) were estimated based on total particle removal (mean FR), and that for each size channel (giving maximum FR). Mean and maximum filtering rates increased from NVI (Nauplius Stage VI) through CVI (Copepodid Stage VI). Both rates plotted against weight satisfied a log fit best for A. tonsa, and a linear fit best for E. affinis. Results for A. tonsa were quite variable, apparently due to differences in temperature between experiments. Particle selection was investigated from the shape of the filtering rate curve over particle size. We define selective feeding by a FR curve which is higher in some size categories, and non-selective feeding by a flat FR curve. The general pattern was one of selective feeding in all copepodid stages of the three calanoid copepods investigated. E. affinis tended to track biomass peaks while Acartia spp.'s feeding was more variable, including feeding in size ranges of greatest particle concentration, on larger particles, and in other size categories as well. Experiments with nauplii tended to yield flat FR curves, and it may be that selective grazing appears with, or is greatly accentuated by, metamorphosis from NVI to CI (Copepodid Stage I).University of Maryland, Center for Environmental and Estuarine Studies Contribution No. 762.     

Etude expérimentale de la ponte chez trois espèces de copépodes pélagiques (Centropages typicus,Acartia clausi et Temora stylifera)

Egg-laying of 3 common copepod species from the Gulf of Marseilles (Centropages typicus Kröyer, Acatia clausi Giesbrecht, Temora stylifera Dana) has been studied under various trophic conditions, at different periods of the year. The role of phytoplankton abundance in the induction and importance of egg-laying is indicated. The specific quality of the algal suspension used for feeding affected also fertility. A seasonal variation in the importance of egg-laying is established. Some observations are made on egg deposition and development.     

Feeding opportunities of larval and juvenile cod (Gadus morhua) in a Greenlandic fjord: temporal and spatial linkages between cod and their preferred prey

Feeding of fish depends on a spatial and temporal match with prey, and since larval and juvenile feeding can be highly selective, their preferences for given prey sizes and taxa should be considered when quantifying the actual availability of potential prey. We investigated the diet and prey preferences of the early-life stages of Atlantic cod (Gadus morhua) to quantify the availability of prey during a spring-summer season in a West Greenlandic fjord. We hypothesized that abundances of larval and juvenile cod at size were synchronized to optimal availability of preferred prey in space and time. The present analysis is based on nine cruises each covering 5 stations visited between 24 May and 5 August 2010 comparing zooplankton abundance, cod gut content and distribution patterns. Cod 4–25 mm in length preferred prey of about 5 % of their own length. During ontogeny, their preferences changed from calanoid nauplii towards Pseudocalanus spp. and Calanus spp. copepodites. The larvae/juvenile had an exceptionally high dietary contribution from cladocerans, which were highly preferred by cod larger than 9 mm, while the abundant Metridia longa and the non-calanoid copepods contributed less. These findings stress the importance of focusing on abundance of preferred prey when assessing the actual prey availability to young fish. We found a spatio-temporal overlap between cod and their preferred prey, and observations suggest that advection of both zooplankton and cod contributed to this overlap. Hence, the larval feeding opportunities might be sensitive to climate-related changes affecting the circulation patterns in this fjord.     

Natural diets of vertically migrating zooplankton in the Sargasso Sea

The feeding preferences of three common diel vertically migrating zooplankton were investigated from December 1999 to October 2000 at the U.S. JGOFS Bermuda Atlantic Time-Series Study (BATS) station in the Sargasso Sea. Gut content analysis of the copepods Pleuromamma xiphias (Giesbrecht) and Euchirella messinensis (Claus) and of the euphausiid Thysanopoda aequalis (Hansen) indicated that all three species fed on a wide variety of phytoplankton, zooplankton, and detrital material. Diet changes generally reflected seasonal trends in phytoplankton community structure. However, species-specific feeding preferences and differences in feeding selectivity among the three species were evident, and in general agreement with feeding habits predicted from the analysis of mouthpart morphology. The euphausiid T. aequalis fed equally on more different food types compared to both copepod species. The copepod P. xiphias consumed a diverse assemblage of phytoplankton from late winter through the summer (78-93% of gut items, by number, were phytoplankton) and based its diet more strongly on carnivorous feeding in autumn and early winter (31% and 61% of gut items were phytoplankton, respectively). E. messinensis showed the greatest feeding specialization, with a strong preference for pennate diatoms in winter and spring and for coccolithophorids during late summer and fall (constituting 67-93% of gut items by number). All three species consumed diatoms more than other phytoplankton taxa, even though diatoms form only a small fraction of the phytoplankton biomass in the Sargasso Sea. Although the majority of gut items identified were phytoplankton cells, the relative biomass contribution of these small cells may be lower than that of zooplankton and detritus. Zooplankton on which the three species primarily preyed were protozoans and crustaceans, but also included other metazoans such as chaetognaths and cnidarians. Marine snow was also an important component of the diet in all three species, with typically >50% and rarely <20% of the gut content being olive-green debris. Marine snow from larvacean houses was found in the guts of all three species, while E. messinenis appeared to selectively consume marine snow aggregates enriched with bicapitate Nitzschia spp. Large cyanobacteria (>4 µm in diameter) found in guts were also likely consumed with marine snow. The species-specific differences in the diets of these three migrating species suggest that an individual species approach is important in determining how feeding habits affect the structure of pelagic food webs and carbon cycling in the sea. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00227-002-0815-8.     

Strain-related physiological and behavioral effects of <Emphasis Type="Italic">Skeletonema marinoi</Emphasis> on three common planktonic copepods

Three strains of the chain-forming diatom Skeletonema marinoi, differing in their production of polyunsaturated aldehydes (PUA) and nutritional food components, were used in experiments on feeding, egg production, hatching success, pellet production, and behavior of three common planktonic copepods: Acartia tonsa, Pseudocalanus elongatus, and Temora longicornis. The three different diatom strains (9B, 1G, and 7J) induced widely different effects on Acartia tonsa physiology, and the 9B strain induced different effects for the three copepods. In contrast, different strains induced no or small alterations in the distribution, swimming behavior, and turning frequency of the copepods. 22:6(n-3) fatty acid (DHA) and sterol content of the diet typically showed a positive effect on either egg production (A. tonsa) or hatching success (P. elongatus), while other measured compounds (PUA, other long-chain polyunsaturated fatty acids) of the algae had no obvious effects. Our results demonstrate that differences between strains of a given diatom species can generate effects on copepod physiology, which are as large as those induced by different algae species or groups. This emphasizes the need to identify the specific characteristics of local diatoms together with the interacting effects of different mineral, biochemical, and toxic compounds and their potential implications on different copepod species.     

Patterns and selectivity in the feeding of certain mesopelagic fishes

Stomach contents were analysed from the 7 most numerous species of mesopelagic fish caught in a series of 11 hauls over a 24 h period at 230 to 266 m depth in the eastern North Atlantic Ocean. The numerical abundance of organisms per filled stomach and the frequency of occurrence of empty stomachs were used to indicate feeding periodicity. The ecological significance of the feeding periodicity was considered by examining it in connection with an investigation of the day-night vertical distribution of zooplankton and micronekton to 2000 m at the same station. Additional dietary evidence on the 7 species considered was also obtained from the vertical series. Feeding selectivity was examined by comparing the composition of the zooplankton population, sampled separately but simultaneously with the micronekton, with that from the overall stomach contents of the species examined. Feeding periodicity was demonstrated for 6 species, of which 3 were found to be feeding selectively: Valenciennellus tripunctulatus on calanoid copepods, Argyropelecus aculeatus on ostracods, and Lampanyctus cuprarius on amphipods and possibly euphausiids. The limited data available on the other 3 species suggested that they were either random feeders (A. hemigymnus and Lobianchia dofleini) or perhaps selecting against a particular group (Notolychnus valdiviae). No indication of feeding periodicity or selectivity was found for Chauliodus danae. The overall pattern of results confirmed the supposed close correlation between vertical migration and feeding in mesopelagic fish.     

Ingestion and incorporation of coral-mucus detritus by reef zooplankton

The copepod Acartia tonsa and the reef mysid Mysidium integrum ingest stained coral mucus. Ingestion rates determined with radioisotope-labeled mucus ranged from 4 to 81% body carbon · 24 h^-1 for the copepods and I to 70% body carbon · 24 h^-1 for the mysids. Incorporation was measured by comparing the organic composition of fecal material and by the incorporation of isotope-labeled mucus. A. tonsa incorporated 47% of ingested ash-free material, 68% of carbon and 36% of nitrogen. M. integrum incorporated 44% of ingested ash-free matter, 57% of carbon and 55% of nitrogen. Incorporation estimates using ¹⁴C-labeled mucus were 65% and 39% for incorporation by A. tonsa and M. integrum respectively. A. tonsa and M. integrum incorporated both the mucus substrate and the epiphytic bacteria of the mucus-detritus.     

Physical and biological controls of algal blooms in the Río de la Plata

Coupled three-dimensional hydrodynamic and ecological numerical simulations were used to investigate the role of transport, stagnation zones and dispersion on inter-annual blooms of the diatom Aulacoseira sp. in the vicinity of the drinking water intakes of the Buenos Aires city (Argentina) in the upper Río de la Plata. Three different summer events were analyzed. First, a mild biomass bloom year (2006–2007), second, a high biomass bloom year (2007–2008) and third, a “normal” no bloom year (2009–2010). Simulated water height, water temperature, suspended solids and chlorophyll \(a\) concentrations patterns compared well with field data. Results revealed that the advection of phytoplankton cells via inflows to the Río de la Plata triggered Aulacoseira sp. blooms in the domain. In addition, excessive growth observed near the drinking water intakes, along the Argentinean margin, were associated with long retention times (stagnant region) and weak horizontal dispersion. Increased concentrations of suspended solids in the water column, in response to re-suspension events, did not prevent the blooms, however, were found to also play a key role in controlling the rate of phytoplankton growth. Finally, a non-dimensional parameter, R, that considers phytoplankton patch size, e-folding growth and dispersion time scales is shown to determine the potential bloom occurrences, as well as bloom intensity; R values higher than 5.7 suggest intense phytoplankton growth. For the mild biomass bloom year, \(R = 7.5\) , for the high biomass bloom year, \(R = 11\) and for the “normal” no bloom year \(R= 0.4\) .