Morphology and sinking velocities of fecal pellets of copepod, molluscan, euphausiid, and salp taxa in the northeastern tropical Atlantic

Morphological characteristics and sinking velocities of naturally occurring fecal pellets of copepods, euphausiids, salps, and pelagic mollusks collected in the northeastern tropical Atlantic were investigated during the period of May-June 1992. The fecal pellets of copepods and euphausiids were cylindrical and distinguished only by their size. Those of salps were, in general, rectangular, and slight differences were noted according to the species. The fecal pellets of the molluscan pteropod Clio sp. were conical, while those of the molluscan heteropod Carinaria sp. were spiral. The sinking velocities ranged from 26.5 to 159.5 m day^-1 for copepod fecal pellets, from 16.1 to 341.1 m day^-1 for euphausiid pellets, from 43.5 to 1167.6 m day^-1 for salps' pellets (Cyclosalpa affinis, Salpa fusiformis, Iasis zonaria, and two unidentified species), from 65 to 205.7 m day^-1 for Clio sp. pellets, and from 120.3 to 646.4 m day^-1 for Carinaria sp. fecal pellets. The measured sinking velocities were compared with estimates predicted using the equations of Komar et al. (1981; Limnol Oceanogr 26:172-180), Stokes' law, and Newton's second law, using either a constant density of fecal pellets (1.22 g cm^-3) or densities estimated with the three different equations. When a constant density was used, the three equations overestimated the sinking velocities; Stokes' law resulted in the largest overestimation, and Newton's second law, the smallest. At the taxa level, the overestimation was greatest for euphausiid 1 fecal pellets and smallest for copepod fecal pellets. When the three equations were used to estimate fecal pellet density, the density estimated using the equation of Komar et al. was the greatest, and that using Stokes' law, the smallest, resulting in over- and underestimation of sinking velocities, respectively. Newton's second law resulted in an intermediate density and gave the closest estimate of sinking velocities. We propose that measurement of sinking velocities of a portion of the fecal pellets might guide in choosing an appropriate equation to be used for a reasonable interpretation of vertical mass flux.     

Are monoalgal diets inferior to plurialgal diets to maximize cultivation of the calanoid copepod Temora stylifera?

Temora stylifera adult copepods were fed with four different monoalgal diets and six combinations of the same cultures for 15 days. Fecundity, hatching success, number of cannibalized embryos, fecal pellet production, adult mortality and naupliar recruitment were compared, in order to find the best diet for this species. Phytoplankton species tested were Prorocentrum minimum (PRO); Isochrysis galbana (ISO); Tetraselmis suecica (TETRA) and Rhodomonas baltica (RHO) which were supplied alone or in different combinations and at various concentrations ranging from a minimum of 1 mg C L⁻¹ day⁻¹ to a maximum of 66 mg C L⁻¹ day⁻¹. Of the ten diets tested, ISO was the worst and was unable to sustain egg production and adult survival possibly because adults were unable to ingest this alga due to its small size. TETRA was also a poor food since it negatively impacted egg production and adult survival, as well as egg hatching success, possibly due to the lack of essential compounds necessary for optimal embryogenesis. RHO and PRO were the best foods inducing highest egg production, hatching success and naupliar recruitment. Even if mean egg production rates were similar to those obtained with some mixed diets, carbon intake concentrations with mixed diets were from 3 to 33 and from 6.6 to 66 times higher than with RHO and PRO given alone, respectively. Mixed diets of ISO and PRO, especially when supplied at higher concentrations (66 mg C L⁻¹ day⁻¹), had a negative effect on egg hatching success and adult survival, with a corresponding reduction in naupliar recruitment. On the other hand, mixed diets of TETRA and PRO promoted high naupliar recruitment but values were similar to PRO offered alone. Our results indicate that a good monoalgal diet such as RHO and PRO can be as effective as a mixed diet to sustain the mass cultivation of T. stylifera.     

Sinking rates of natural copepod fecal pellets

Many pure samples of natural fecal pellets have been collected from mixed small copepods and from the pontellid copepod Anomalocera patersoni in the Ligurian Sea, using a specially designed pellet collection device. Sinking rates of fresh pellets and pellets aged up to 33 days have been determined at 14°C, the mean temperature of the essentially isothermal water column in the Ligurian Sea. Sinking rates of pellets collected during calm sea states increased with increasing pellet volume, but sinking rates of pellets collected during rough sea (Beaufort scale 6) showed little correlation with pellet size. Much of the variability in the sinking rate-pellet size relationships was the result of different pellet composition and compaction, but not pellet age. Pellets produced from laboratory diets of phytoplankton and phytoplankton-sediment mixes showed the expected wide variability in sinking rates, with sediment-ballasted pellets sinking much faster than pellets produced from pure algal diets; thus determination of vertical material fluxes in the sea using laboratory-derived fecal pellet sinking rates is unwarranted. Natural pellet sinking data for small copepods and A. patersoni have been combined with similar data for euphausiids, to yield sinking rates of roughly two orders of magnitude over three orders of magnitude in pellet volume. Pellets from small copepods sank at speeds too slow to be of much consequence to rapid material flux to the deep sea, but they undoubtedly help determine upper water distribution of materials. Recalculation of fecal pellet mass flux estimates from the literature, using our sinking rate data for natural small copepod pellets, yielded estimates about half those of previously published values. Earlier studies had concluded that small fecal pellets were of lesser significance to total material flux than fecal matter; our recalculation strengthens that conclusion. Pellets from large copepods and euphausiids, however, have the capability to transport materials to great depths, and probably do not substantially recycle materials near the surface. The fact that the majority of pellets which had previously been collected in deep traps by other workers were of a size comparable to pellets from our large copepods supports the contention that these larger pellets are the main ones involved in vertical flux.     

Defecation by Salpa thompsoni and its contribution to vertical flux in the Southern Ocean

Measurements of the defecation rate of Salpa thompsoni were made at several stations during two cruises west of the Antarctic Peninsula in 2004 and 2006. Rates were quantified in terms of number of pellets, pigment, carbon and nitrogen for a wide size range of both aggregate and solitary salps. Measured defecation rates were constant over several hours when salps were held at near-surface conditions from which they had been collected. The defecation rate per salp increased with both salp size and the ambient level of particulate organic matter (POM) in the upper water column. The weight-specific defecation rate ranged between 0.5 and 6% day⁻¹ of salp body carbon, depending on the concentration of available particulate matter in the water. Carbon defecation rates were applied to biomass estimates of S. thompsoni to calculate daily carbon defecation rates for the populations sampled during the two cruises. Dense salp populations of over 400 mg C m⁻² were calculated to produce about 20 mg C m⁻² day⁻¹, comparable to other major sources of vertical flux of organic material in the Southern Ocean. Measured sinking rates for salp fecal pellets indicated that the majority of this organic material could reach deep sediments within a few days, providing a fast and direct pathway for carbon to the deep ocean.     

Summer copepod production in subtropical waters adjacent to Australia's North West Cape

Growth and secondary production of pelagic copepods near Australia's North West Cape (21° 49 S, 114° 14 E) were measured during the austral summers of 1997/1998 and 1998/1999. Plankton communities were diverse, and dominated by copepods. To estimate copepod growth rates, we incubated artificial cohorts allocated to four morphotypes, comprising naupliar and copepodite stages of small calanoid and oithonid copepods. Growth rates ranging between 0.11 and 0.83 day^–1 were low, considering the high ambient temperatures (23–28°C). Calanoid nauplii had a mean growth rate of 0.43±0.17 day^-1 (SD) and calanoid copepodites of 0.38±0.13 day^-1. Growth rates of oithonid nauplii and copepodites were marginally less (0.38±0.19 day^–1 and 0.28±0.11 day^–1 respectively). The observed growth rates were suggestive of severe food limitation. Although nauplii vastly outnumbered copepodite and adult copepods, copepodites comprised the most biomass. Copepodites also contributed most to secondary production, although adult egg production was sporadically important. The highest copepod production was recorded on the shelf break (60 mg C m^-2 day^-1). Mean secondary production over both shelf and shelf break stations was 12.6 mg C m^-2 day^-1. Annual copepod secondary production, assuming little seasonality, was estimated as ~ 3.4 g C m^-2 year^-1 (182 kJ m^-2 year^-1).Communicated by G.F. Humphrey, Sydney     

Seasonal variations in the densities of fecal pellets produced by Oikopleura vanhoeffeni (C. Larvacea) and Calanus finmarchicus (C. Copepoda)

Two abundant macrozooplankters, Oikopleura vanhoeffeni (Lohmann) and Calanus finmarchicus (Gunnerus) were collected from the coastal waters off Newfoundland in different seasons during 1990–1991 and incubated in natural seawater to collect freshly egested, field produced, fecal pellets. The densities of fecal pellets from O. vanhoeffeni and C. finmarchicus were measured in an isosmotic density gradient. These are the first reported seasonal measurements of fecal pellet densities from two different types of macrozooplankters, O. vanhoeffeni, a larvacean, filter feeder and C. finmarchicus, a crustacean, suspension feeder. Pellet density ranges and medians were significantly different among seasons for both species, depending primarily on the type of phytoplankton ingested and its ability to be compacted. Winter O. vanhoeffeni and fall C. finmarchicus feces filled with nanoplankters and soft bodied organisms had less open space [packing index (% open area) = 3.5 and 4% for O. vanhoeffeni and C. finmarchicus, respectively] and were more dense (1.23 and 1.19 g cm^-3) than spring feces filled with diatoms (packing index = 15 and 23%, density = 1.13 and 1.11 gcm^-3). For copepods, these results contrast with previously published density values and with the predicted copepod fecal pellet density calculated, in the present study, using the conventional mass/volume relationship. Copepod spring and summer diatom-filled feces had a calculated density of 1.12 and 1.24 gcm^-3 vs a measured median density of 1.11 gcm^-3 for both spring and summer feces; the fall feces containing nanoplankters had a calculated density of 1.08 gcm^-3 vs a measured median density of 1.19 gcm^-3. Knowledge of the seasonal variations in fecal pellet densities is important for the development of flux models.     

Comparison of the chemical composition of particulate material and copepod faecal pellets at stations off the coast of Labrador and in the Gulf of St. Lawrence

Faecal pellets were collected in 1988 from copepods which had fed in situ or in laboratory experiments, using screened natural seawater as food, at two stations off the coast of Labrador and one in the Gulf of St. Lawrence. The chemical composition of the pellets and of particulate material in profiles and in laboratory food were measured in terms of particulate carbon, carbohydrate (soluble and insoluble), protein and lipid. Faecal pellet composition was somewhat similar in all experiments at the first two stations, where the compositions of particulate material in situ and copepod species assemblages were also similar. At the third station the compositions of faecal pellets and particulate material were slightly different from those at the other stations and the copepod species composition varied between sampling times. Faecal pellets at the first two stations had very low levels of soluble carbohydrate, while concentrations in the food were generally high, suggesting that it was efficiently metabolized by copepods, although it might have been absent because of sloppy feeding or release, after passage through the gut, in soluble form or from faecal pellets. Comparisons of POC: biogenic silica ratios in food and faecal pellets, calculated using data presented elsewhere (Head 1992; Mar. Biol. 112: 583–592), suggested that at these stations, where food concentrations were high (chlorophyll concentrations>8 gl^-1), copepods may have been assimilating carbon rather inefficiently.     

Life cycle of <Emphasis Type="Italic">Oithona</Emphasis> <Emphasis Type="Italic">similis</Emphasis> (Copepoda: Cyclopoida) in Kola Bay (Barents Sea)

The annual population dynamics (nauplii, old copepodites CIV–CV and adults) and seasonal variations in reproductive parameters of the cyclopoid copepod Oithona similis were investigated on the basis of the data 1999–2006 in Kola Bay, a large subarctic fjord in the Barents Sea. Population density of O. similis ranged from 110 to 9,630 ind m⁻³ and averaged 1,020 ± 336 ind m⁻³. The relative abundance of adults was high during winter (~60%). At the end of winter (mid-March), the population included a large percentage of later-stage copepodites (stage CIV 23% and stage CV 57%). There were two periods of mass spawning, in late June and September. Autumn and summer generations strongly differed in abundance, average prosome length (PL), clutch size (CS), egg diameter (D), egg production rates (EPR and SEPR) and female secondary production. Average PL decreased with increasing water temperature, while D and CS were strongly correlated with PL but unaffected by temperature. Annual average EPR and SEPR were 0.55 ± 0.18 eggs female⁻¹ day⁻¹ and 0.0011 ± 0.003 day⁻¹, respectively. Female secondary production averaged 0.8 ± 0.3 μg C m⁻³ day⁻¹ (range 0.001–3.58). There were positive relationships between abundance, EPR, SEPR, production and water temperatures. Reproductive parameters appeared to be controlled by hydrological factors and food conditions.     

Trophodynamic function of copepods,appendicularians and protozooplankton in the late summer zooplankton community in the Skagerrak

The study was carried out in the Skagerrak during late summer when population development in the pelagic cycle culminated in the yearly maximum in zooplankton biomass. The cyclonic circulation of surface water masses created the characteristic dome-shaped pycnocline across the Skagerrak. The large dinoflagellate Ceratium furca dominated the phytoplankton biomass. Ciliates and heterotrophic dinoflagellates were the major grazers and, potentially, consumed 43–166% of daily primary production. The grazing impact of copepods was estimated from specific egg production rates and grazing experiments. The degree of herbivory differed between species (14–85%), but coprophagy (e.g. feeding on fecal pellets) and ingestion of microzooplankton were also important. The appendicularian Oikopleura dioica was present in lower numbers than copepods, but cleared a large volume of water. The grazing impact of copepods and O. dioica was estimated to 57±24% and 12±12% of daily primary production, respectively. Sedimentation of organic material (30 m) varied between 169 and 708 mg C m^–2 day^–1, and the contribution from the mesozooplankton (copepod fecal pellets and mucus houses with attached phytodetritus of O. dioica) was 5–33% of this sedimentation. Recycling of fecal pellets and mucus houses in the euphotic zone was 59% and 36%, respectively. However, there was a high respiration of organic material by microorganisms in the mid-water column, and 34% of the sedimenting material actually reached the benthic community in the deep, central part of the Skagerrak.     

The tunicate Salpa thompsoni ecology in the Southern Ocean. I. Distribution, biomass, demography and feeding ecophysiology

Distribution, density, and feeding dynamics of the pelagic tunicate Salpa thompsoni have been investigated during the expedition ANTARKTIS XVIII/5b to the Eastern Bellingshausen Sea on board RV Polarstern in April 2001. This expedition was the German contribution to the field campaign of the Southern Ocean Global Ocean Ecosystems Dynamics Study (SO-GLOBEC). Salps were found at 31% of all RMT-8 and Bongo stations. Their densities in the RMT-8 samples were low and did not exceed 4.8 ind m⁻² and 7.4 mg C m⁻². However, maximum salp densities sampled with the Bongo net reached 56 ind m⁻² and 341 mg C m⁻². A bimodal salp length frequency distribution was recorded over the shelf, and suggested two recent budding events. This was also confirmed by the developmental stage composition of solitary forms. Ingestion rates of aggregate forms increased from 2.8 to 13.9 μg (pig) ind⁻¹ day⁻¹ or from 0.25 to 2.38 mg C ind⁻¹ day⁻¹ in salps from 10 to 40 mm oral-atrial length, accounting for 25–75% of body carbon per day. Faecal pellet production rates were on average 0.08 pellet ind⁻¹ h⁻¹ with a pronounced diel pattern. Daily individual egestion rates in 13 and 30 mm aggregates ranged from 0.6 to 4.8 μg (pig) day⁻¹ or from 164 to 239 μg C day⁻¹. Assimilation efficiency ranged from 73 to 90% and from 65 to 76% in 13 and 30 mm aggregates, respectively. S. thompsoni exhibited similar ingestion and egestion rates previously estimated for low Antarctic (~50°S) habitats. It has been suggested that the salp population was able to develop in the Eastern Bellingshausen Sea due to an intrusion into the area of the warm Upper Circumpolar Deep Water     

Phytoplankton growth and microzooplankton grazing rates in a restricted Mediterranean lagoon (Bizerte Lagoon,Tunisia)

Phytoplankton growth and microzooplankton grazing were investigated in the restricted Bizerte Lagoon in 2002 and 2004. The 2002 study, carried out at one station from January to October, showed significant seasonal variations in phytoplankton dynamics. High growth rates (0.9–1.04 day⁻¹), chlorophyll a (Chl a) concentrations (6.6–6.8 μg l⁻¹) and carbon biomass (392–398 μg C l⁻¹) were recorded in summer (July), when several chain-forming diatoms had intensively proliferated and dominated the carbon biomass (74%). In 2004, four stations were studied during July, a period also characterized by the high proliferation of several diatoms that made up 70% of the algal carbon biomass. In 2004, growth rates (0.34–0.45 day⁻¹) and biomass of algae (2.9–5.4 μg Chl a l⁻¹ and 209–260 μg C l⁻¹) were low, which may be related to the lower nutrient concentrations recorded in 2004. Microzooplankton >5 μm were mainly composed of heterotrophic dinoflagellates and ciliates. Microzooplankton biomass peaked during summer (2002 320–329, 2004 246–361 μg C l⁻¹), in response to the enhanced phytoplankton biomass and production. The grazer biomass was dominated by ciliates (71–76%) in July 2002 and by heterotrophic dinoflagellates (52–67%) in July 2004. Throughout the year and at different stations, microzooplankton grazed actively on phytoplankton, removing 26–58% of the Chl a and 57-84% of the primary production. In 2002, the highest grazing impact was observed on the large algae (>10 μm) during the period of diatom dominance. These results have a significant implication for carbon export to depth. Indeed, the recycling of most of the diatom production by the microbial food web in the upper water column would reduce the flux of material to the seafloor. This should be considered when modeling the carbon cycling in coastal environments and under conditions of diatom dominance. During both studies, ciliates had higher growth rates (0.5–1.5 day⁻¹) and a higher carbon demand (165–470 μg C l⁻¹ day⁻¹) than dinoflagellates (0.1–0.5 day⁻¹, 33–290 μg C l⁻¹ day⁻¹). Moreover, when grazer biomass was dominated by ciliates (in July 2002), herbivory accounted for 71–80% of the C ingested by microzooplankton while it accounted only for 14–23% when dinoflagellates dominated the grazer biomass (in July 2004). These results suggest that, in contrast to findings from open coastal waters, ciliate species of the restricted Bizerte Lagoon were more vigorous grazers of the large algae (diatoms) than were dinoflagellates.     

Diet and egg production of the copepod Acartia tonsa in Florida Bay. II. Role of the nutritional environment

 As part of an ongoing study of changes in the trophic pathways of Florida Bay's pelagic ecosystem, the nutritional environment (seston protein, lipid and carbohydrate levels), diet (taxon-specific microplankton ingestion rates) and egg production rate of the important planktonic copepod Acartia tonsa were measured off Rankin and Duck Keys in July and September 1997 and in January, March and May 1998. Rankin Key has been the site of extensive sea grass mortality and persistent ultraplankton blooms since 1987. Duck Key has experienced neither of these perturbations. Protist (auto-plus heterotroph) biomass was approximately twice as high off Rankin as off Duck Key. Diatoms, dinoflagellates and heterotrophic protists dominated the food environment off Rankin Key, while cells <5 μm diam often predominated off Duck Key. Protein and carbohydrate concentrations were higher off Rankin Key than Duck Key, while average lipid levels were usually low at both stations. Ingestion rates at both stations frequently approached temperature- and food-dependent maxima for the species, exceeding 100% of estimated body C d⁻¹ on 3 of 5 occasions off Rankin Key. Egg production rates, however, were consistently low (Rankin: 3 to 16 eggs copepod⁻¹ d⁻¹; Duck: 1 to 12 eggs copepod⁻¹ d⁻¹), and gross egg production efficiencies (100% × egg production C/ingested C) averaged <10%. At Duck Key, egg production rate varied with temperature and food concentration, while off Rankin Key, egg production was strongly correlated with seston protein content. The efficiency with which lipids (which were scarce in the seston) were transferred from the diet to the eggs increased exponentially with decreasing seston lipid content. Egg production efficiencies based on protein, however, were independent of seston protein content and never exceeded 10%. Received: 23 December 1998 / Accepted: 23 March 2000     

Microbial activity and carbon, nitrogen, and phosphorus content in a subtropical seagrass estuary (Florida Bay): evidence for limited bacterial use of seagrass production

Bacterial abundance, production, and extracellular enzyme activity were determined in the shallow water column, in the epiphytic community of Thalassia testudinum, and at the sediment surface along with total carbon, nitrogen, and phosphorus in Florida Bay, a subtropical seagrass estuary. Data were statistically reduced by principle components analysis (PCA) and multidimensional scaling and related to T. testudinum leaf total phosphorus content and phytoplankton biomass. Each zone (i.e., pelagic, epiphytic, and surface sediment community) was significantly dissimilar to each other (Global R = 0.65). Pelagic aminopeptidase and sum of carbon hydrolytic enzyme (esterase, peptidase, and α- and β-glucosidase) activities ranged from 8 to 284 mg N m⁻² day⁻¹ and 113–1,671 mg C m⁻² day⁻¹, respectively, and were 1–3 orders of magnitude higher than epiphytic and sediment surface activities. Due to the phosphorus-limited nature of Florida Bay, alkaline phosphatase activity was similar between pelagic (51–710 mg P m⁻² day⁻¹) and sediment (77–224 mg P m⁻² day⁻¹) zones but lower in the epiphytes (1.1–5.2 mg P m⁻² day⁻¹). Total (and/or organic) C (111–311 g C m⁻²), N (9.4–27.2 g N m⁻²), and P (212–1,623 mg P m⁻²) content were the highest in the sediment surface and typically the lowest in the seagrass epiphytes, ranging from 0.6 to 8.7 g C m⁻², 0.02–0.99 g N m⁻², and 0.5–43.5 mg P m⁻². Unlike nutrient content and enzyme activities, bacterial production was highest in the epiphytes (8.0–235.1 mg C m⁻² day⁻¹) and sediment surface (11.5–233.2 mg C m⁻² day⁻¹) and low in the water column (1.6–85.6 mg C m⁻² day⁻¹). At an assumed 50% bacterial growth efficiency, for example, extracellular enzyme hydrolysis could supply 1.8 and 69% of epiphytic and sediment bacteria carbon demand, respectively, while pelagic bacteria could fulfill their carbon demand completely by enzyme-hydrolyzable organic matter. Similarly, previously measured T. testudinum extracellular photosynthetic carbon exudation rates could not satisfy epiphytic and sediment surface bacterial carbon demand, suggesting that epiphytic algae and microphytobenthos might provide usable substrates to support high benthic bacterial production rates. PCA revealed that T. testudinum nutrient content was related positively to epiphytic nutrient content and carbon hydrolase activity in the sediment, but unrelated to pelagic variables. Phytoplankton biomass correlated positively with all pelagic components and sediment aminopeptidase activity but negatively with epiphytic alkaline phosphatase activity. In conclusion, seagrass production and nutrient content was unrelated to pelagic bacteria activity, but did influence extracellular enzyme hydrolysis at the sediment surface and in the epiphytes. This study suggests that seagrass-derived organic matter is of secondary importance in Florida Bay and that bacteria rely primarily on algal/cyanobacteria production. Pelagic bacteria seem coupled to phytoplankton, while the benthic community appears supported by epiphytic and/or microphytobenthos production.     

Feeding and predation impact of two chaetognath species, Eukrohnia hamata and Sagitta gazellae, in the vicinity of Marion Island (southern ocean)

The predation impact of the two chaetognaths Eukrohnia hamata and Sagitta gazellae on mesozooplankton standing stock were investigated in three depth layers during two 24 h stations occupied in the vicinity of Marion Island in late austral summer (April/May) 1986. The zooplankton community at both stations was dominated by small copepods (Oithona spp., Microcalanus spp.), which accounted for >95% of total zooplankton abundance. Chaetognaths comprised <2% of total zooplankton abundance. E. hamata constituted >95% of the total chaetognath stock. The general trend in both species was decreasing abundance with increasing depth, which appeared to be correlated to the distribution of copepods (r ² = 0.45; P <0.05). Gut-content analysis showed that copepods (mainly Oithona spp., Calanus spp. and Rhincalanus gigas) and ostracods were the main prey of both species, accounting for 87 and 61% of the total number of prey in E. hamata and S.␣gazellae stomachs, respectively. In the guts of S.␣gazellae, pteropods (Limacina spp.) and chaetognaths were also well represented. The mean number of prey items (NPC) for E. hamata ranged from 0.02 to 0.06 prey individual⁻¹ which corresponds to an individual feeding rate (Fr) of between 0.05 and 0.12 prey d⁻¹. For S.␣gazellae, the NPC values were higher, varying between 0.04␣and 0.20 prey individual⁻¹, or between 0.15 and 0.76 prey d⁻¹. The daily predation impact of the two chaetognaths was estimated at between 0.3 and 1.2% of the copepod standing stock or between 7 and 16% of the daily copepod production. Predation by S. gazellae on chaetognaths accounted for up to 1.6% of the chaetognath standing stock per day. Received: 26 November 1996 / Accepted: 31 October 1997     

Prey, feeding rates, and asexual reproduction rates of the introduced oligohaline hydrozoan Moerisia lyonsi

Moerisia lyonsi Boulenger (Hydrozoa) medusae and benthic polyps were found at 0 to 5‰ salinity in the Choptank River subestuary of Chesapeake Bay, USA. This species was introduced to the bay at least 30 years before 1996. Medusae and polyps of M. lyonsi are very small and inconspicuous, and may occur widely, but unnoticed, in oligohaline waters of the Chesapeake Bay system and in other estuaries. Medusae consumed copepod nauplii and adults, but not barnacle nauplii, polychaete and ctenophore larvae or tintinnids, in laboratory experiments. Predation rates on copepods by medusae increased with increasing medusa diameter and prey densities. Feeding rates on copepod nauplii were higher than on adults and showed no saturation over the range of prey densities tested (1 to 64 prey l⁻¹). By contrast, predation on copepod adults was maximum (1 copepod medusa⁻¹ h⁻¹) at 32 and 64 copepods l⁻¹. Unexpectedly, M. lyonsi colonized mesocosms at the Horn Point Laboratory during the spring and summer in 4 years (1994 to 1997), and reached extremely high densities (up to 13.6 medusae l⁻¹). Densities of copepod adults and nauplii were low when medusa densities were high, and estimated predation effects suggested that M. lyonsi predation limited copepod populations in the mesocosms. Polyps of M. lyonsi asexually produced both polyp buds and medusae. Rates of asexual reproduction increased with increasing prey availability, from an average total during a 38 d experiment of 9.5 buds polyp⁻¹ when each polyp was fed 1 copepod d⁻¹, to an average total of 146.7 buds polyp⁻¹ when fed 8 copepods d⁻¹. The maximum daily production measured was 8 polyp buds and 22 medusae polyp⁻¹. The colonizing potential of this hydrozoan is great, given the high rates of asexual reproduction, fairly wide salinity tolerance, and existence of a cyst stage. Received: 29 October 1998 / Accepted: 3 March 1999     

Production of tropical copepods in Kingston Harbour, Jamaica: the importance of small species

The copepod community observed during an 18-month period at the mouth of eutrophic Kingston Harbour, Jamaica, was dominated by small species of Parvocalanus, Temora, Oithona, and Corycaeus. Mean copepod biomass was 22.1 mg AFDW m⁻³ (331 mg m⁻²). Annual production was 1679 kJ m⁻², partitioned as 174 kJ m⁻² naupliar, 936 kJ m⁻² copepodite, 475 kJ m⁻² egg and 93 kJ m⁻² exuvial production. All nauplii, most copepodites and many adults, equivalent to half of the biomass and production, were missed by a standard 200-μm plankton net, emphasizing the importance of nauplii and small species in secondary production estimates. The evidence suggests that growth rates and production are generally not food limited, and we speculate that size-selective predation shapes the structure of the harbour community. Biomass and production are higher than previous estimates for tropical coastal waters, but comparable to other eutrophic tropical embayments and many productive temperate ecosystems. Far from being regions of low productivity, tropical zooplankton communities may have significant production and deserve greater research attention than they currently receive. Received: 19 September 1997 / Accepted: 21 October 1997     

Scanning electron microscopy investigation of bacterial colonization of the marine copepod Acartia clausi

Association between copepods and bacteria was observed in many scanning electron micrographs. Particular sites on the copepods were selectively colonized by bacteria; the joints of segments and legs, swimming legs and depressed parts of the body surface were found to be densely covered with bacteria. In comparison, bacterial attachment to copepod skeletons in fecal pellets excreted by chaetognaths was not selective; bacteria were sparsely found all over the copepod. Between 9 to 30% of copepods in Tokyo Bay waters had attached bacteria in January and April 1983.     

Life cycle of <Emphasis Type="Italic">Pseudocalanus acuspes</Emphasis> Giesbrecht (Copepoda,Calanoida) in the Central Baltic Sea: II. Reproduction,growth and secondary production

The population dynamics of Pseudocalanus acuspes in the Central Baltic Sea were studied from March 2002 to May 2003 on a monthly basis. All stages were present year round with a stage shift from nauplii to older copepodite stages over the course of the year. Biomass, estimated from prosome length, peaked between May and September with maximum recorded values of 594 and 855 mg C m⁻² in May 2002 and 2003, respectively. Differences in biomass between stations up to a factor of 20 were observed especially in April/May and October. Mean egg production rate (EPR) showed a seasonal course and was highest in April 2002 and 2003 with 3.6 and 2.1 eggs f⁻¹ day⁻¹, respectively, corresponding to a mean weight-specific egg production rate (SEPR) of 0.13 and 0.04. Egg production seems to be limited by food from May on. Stage durations determined from moulting experiments turned out to be extremely long. Maximum growth rates based on stage durations of 15–25 days at 4°C in May and July 2003 amounted for 0.03–0.05 day⁻¹ in CI-CIV. Comparing these rates with rates derived from temperature–development relationships for P. acuspes from the literature resulted in five times higher growth rates for the latter case. Secondary production reached values up to 9.1 mg C m⁻² day⁻¹ (method for continuously reproducing populations) and 10.5 mg C m⁻² day⁻¹ (increment summation).     

In situ net primary productivity and photosynthesis of Antarctic sea ice algal, phytoplankton and benthic algal communities

Primary production at Antarctic coastal sites is contributed from sea ice algae, phytoplankton and benthic algae. Oxygen microelectrodes were used to estimate sea ice and benthic primary production at several sites around Casey, a coastal area in eastern Antarctica. Maximum oxygen export from sea ice was 0.95 mmol O₂ m⁻² h⁻¹ (~11.7 mg C m⁻² h⁻¹) while from the sediment it was 6.08 mmol O₂ m⁻² h⁻¹ (~70.8 mg C m⁻² h⁻¹). When the ice was present O₂ export from the benthos was either low or negative. Sea ice algae assimilation rates were up to 3.77 mg C (mg Chl-a)⁻¹ h⁻¹ while those from the benthos were up to 1.53 mg C (mg Chl-a)⁻¹ h⁻¹. The contribution of the major components of primary productivity was assessed using fluorometric techniques. When the ice was present approximately 55–65% of total daily primary production occurred in the sea ice with the remainder unequally partitioned between the sediment and the water column. When the ice was absent, the benthos contributed nearly 90% of the primary production.     

Population dynamics of Eudendrium racemosum (Cnidaria, Hydrozoa) from the North Adriatic Sea

Benthic suspension feeders in shallow waters develop in relation to the food availability and the variation of physical parameters giving rise to complex communities that act as a control factor on the plankton biomass. The aim of the work is to establish the role of the hydrozoan Eudendrium racemosum in the energy transfer from the plankton to the benthos in marine food chains of the North Adriatic Sea. This study highlighted that the hydroid biomass changed over time in relation to temperature and irradiance, and the highest abundance was observed during summer with about 400,000 polyps m⁻² (about 19 g C m⁻²). The population suffered an evident summer decrease in relation to a peak of abundance of its predator, the nudibranch Cratena peregrina, whose adult specimens were able to eat up to 500 polyps day⁻¹ and reached an abundance of 10 individuals m⁻². The gut content analysis revealed that the hydroid diet was based on larvae of other benthic animals, especially bivalves and that the amount of ingested preys changed during the year with a peak in summer when it was estimated an average predation rate of 13.7 mg C m⁻² day⁻¹. In July, bivalves represented over 60 % of the captured items and about 18 mg C m⁻² day⁻¹. Values of biomass of E. racemosum are the highest ever recorded in the Mediterranean Sea, probably supported by the eutrophic conditions of the North Adriatic Sea. Moreover, our data suggest that settling bivalves provide the greater part of the energetic demand of E. racemosum.