Copepod Production Research Articles

Role of midwater chaetognaths in Southern Ocean pelagic energy flow

We estimate the energy flow through mesoand bathypelagic chaetognaths in the Atlantic sector of the Southern Ocean from (1) depth-structured chaetognath abundance and body mass data, (2) a general chaetognath respiration model driven by body mass, temperature, water depth and taxon, and (3) published relationships between respiration, production and consumption in chaetognaths. In the 500 to 2000 m depth layer chaetognaths have a mean biomass of 0.109 mg C m–3 in summer and 0.146 mg C m–3 in winter. Chaetognaths respiration and consumption amount to 282 and 563 mg C m–2 yr–1, respectively. Thus, Antarctic midwater chaetognaths consume 0.05% of the copepod standing stock per day or 1% of the daily copepod production in summer. About 2.8% (= 1.9 g C m–2 yr–1) of the net annual primary production is required to fuel the midwater chaetognath community via herbivorous copepods. When assuming a 1:1 diet of herbivorous and carnivorous copepods, this share increases to 6.1% (= 4.1 g C m–2 yr–1) of annual primary production. It is estimated that chaetognath consumption for the whole water column is 1.4 g C m–2 yr–1. This corresponds to 7.1% (= 4.8 g C m–2 yr–1) and 15.5% (= 10.4 g C m–2 yr–1) of the primary production channeled through herbivorous copepods and through herbivorous and carnivorous copepods, respectively. Hence, chaetognaths represent an important link between lower and higher trophic levels. To further unravel their role in the ecosystem, additional studies on the mesoand bathypelagic zooplankton community are needed.

Explaining regional variability in copepod recruitment: Implications for a changing climate

Characterizing environmental effects on copepod production and their ecological roles is complicated by multiple physical (e.g. temperature) and biological (e.g. food, predation) factors controlling multiple aspects of copepod physiology and demography. For example, data for two regions in eastern Canada (St. John’s, Newfoundland and Halifax, Nova Scotia) indicate that subtle differences in environmental conditions lead to significant differences in seasonal copepod ( Calanus finmarchicus) recruitment timing and magnitude. Here, we quantify how environmental variability influences C. finmarchicus physiology and demography leading to observed regional and seasonal variations in abundance off St. John’s and Halifax. We apply a stochastic individual-based model (IBM) for copepod population dynamics to simulate the seasonal variation in C. finmarchicus abundance of egg through copepodite 1 (C1) stages at the two sites using year-specific local forcing from multi-year data. The model includes individual variability in development, egg production and mortality rates with resulting seasonal C1 abundance averaged among years and compared to analogous observations. We find temperature has a dominant effect on both development and egg production rates while egg recruitment is affected by temperature and female abundance at both sites. We show that mortality rate characterization has a strong influence on modeled abundances, and site-specific environmentally dependent mortality rates are necessary to produce results consistent with observations (temperature vs. food vs. cannibalism via females). Results indicate that prediction of climate change effects on copepod abundance and their ecological roles requires consideration of biological (e.g. chlorophyll a, female abundance) as well as physical (e.g. temperature) factors. In particular, estimates of abundances during the onset of C1 recruitment (i.e. their arrival on the larval fish prey field) are improved by 67–94% when the influence of biological factors on mortality rates are considered.

Role of ciliates and other microzooplankton in the Irminger Sea (NW Atlantic Ocean)

This study focuses on a large region of the open ocean where we predict microzooplankton significantly influence foodweb structure over much of the year. The Irminger Sea exhibits low primary production that is generally poor for copepod production; in such waters ciliates and other microzooplankton can contribute significantly to the diets of holo- and mero- mesozooplankton and are major grazers of primary production. Surface plankton samples were collected during an extensive survey, across the basin and along one transect at several depths, over three seasons (winter, spring, summer), but not including the spring bloom. Microzooplankton and phytoplankton samples were fixed with Lugol’s solution and microscopically enumerated for species abundance; biomass was determined from cell volumes. Basin-scale distributions of abundance, biomass, and production were examined by geostatistical and multidimensional scaling methods. The dominance of the < 10 µm phytoplankton suggests that this should be a microzooplankton-dominated food web. Ciliates and heterotrophic dinoflagellates are abundant, in terms of numbers and biomass; heterotrophic dinoflagellates are more abundant than ciliates, but are less dominant in terms of biomass. Using ciliates as a proxy for all microzooplankton we suggest that there are seasonal patterns in occurrence, and there is no basin-scale patchiness related to hydrographic features. Through some simple, albeit “rough” calculations, we suggest that ciliate production may be sufficient to account for the removal of 15-30% of the < 10 µm primary production. If heterotrophic dinoflagellates were included in these estimates, they may be doubled (i.e. 30-60%). We, thus, contend that microzooplankton are major phytoplankton consumers in the system and should be carefully parameterised in models of this region.

Biological processes in the North Sea: vertical distribution and reproduction of neritic copepods in relation to environmental factors

We studied the vertical distribution and reproduction of dominant neritic copepod species in the Dogger Bank area and surrounding North Sea to reveal (i) if these species are concentrated in the subsurface chlorophyll maximum layer, (ii) if the chlorophyll maximum offers superior food conditions for reproduction compared with surface waters and (iii) if the secondary production is thus higher in the frontal areas with a subsurface chlorophyll maximum. In addition, we wanted to (iv) identify the most important environmental factors determining the reproduction of neritic copepods in the North Sea. We observed a higher egg production of cultured Acartia tonsa when fed with the seston from chlorophyll maximum, but no evidence of a higher copepod abundance in this layer. Secondary production was highest at the station closest to the upwelling of new nutrients, although seasonal differences in environmental variables probably overrode the differences between frontal and stratified stations. Copepod egg production on an annual basis seemed to be best predicted by the body size and specific fatty acids, with a high egg production, but low hatching success associated with a high EPA:DHA ratio. Total secondary production of small copepods seemed mainly related to the species composition, suggesting that factors controlling abundance of specific species rather than reproduction might be more important in determining the secondary production of copepods.

Shearwater Foraging in the Southern Ocean: The Roles of Prey Availability and Winds

BackgroundSooty (Puffinus griseus) and short-tailed (P. tenuirostris) shearwaters are abundant seabirds that range widely across global oceans. Understanding the foraging ecology of these species in the Southern Ocean is important for monitoring and ecosystem conservation and management.Methodology/Principal FindingsTracking data from sooty and short-tailed shearwaters from three regions of New Zealand and Australia were combined with at-sea observations of shearwaters in the Southern Ocean, physical oceanography, near-surface copepod distributions, pelagic trawl data, and synoptic near-surface winds. Shearwaters from all three regions foraged in the Polar Front zone, and showed particular overlap in the region around 140°E. Short-tailed shearwaters from South Australia also foraged in Antarctic waters south of the Polar Front. The spatial distribution of shearwater foraging effort in the Polar Front zone was matched by patterns in large-scale upwelling, primary production, and abundances of copepods and myctophid fish. Oceanic winds were found to be broad determinants of foraging distribution, and of the flight paths taken by the birds on long foraging trips to Antarctic waters.Conclusions/SignificanceThe shearwaters displayed foraging site fidelity and overlap of foraging habitat between species and populations that may enhance their utility as indicators of Southern Ocean ecosystems. The results highlight the importance of upwellings due to interactions of the Antarctic Circumpolar Current with large-scale bottom topography, and the corresponding localised increases in the productivity of the Polar Front ecosystem.

Population structure, egg production and gut content pigment of large grazing copepods during the spring phytoplankton bloom in the Oyashio region

As a basis for analyzing development of six large grazing copepods ( Eucalanus bungii, Metridia pacifica, M. okhotensis, Neocalanus cristatus, N. flemingeri and N. plumchrus) in the Oyashio region, quasi-daily twin-NORPAC net (0.33 and 0.10 mm mesh) hauls were taken through the upper 150 m and 500 m at a station southwest of Hokkaido before (9–14 March) and after (6–30 April) the onset of the phytoplankton bloom in 2007. Based on additional fresh specimens collected from 0–150 m, egg production of E. bungii, M. pacifica and M. okhotensis, and gut pigments of late copepodid stages in each species were evaluated. Total zooplankton biomass was greater from 10 April onward by a factor of 2- to 8-fold the previous levels. This increase of the 0–150 m biomass was caused by development of Neocalanus spp. copepodids and upward migration of resting E. bungii. Egg production of E. bungii peaked on 18 April, while abundance of its nauplii and C1 peaked on 20 and 25 April, respectively. Sex ratio and C6-female gonad maturation index of E. bungii showed new recruitment to C6 during 20–30 April, likely derived from a population that over-wintered as C3 or C4. Egg production and hatchability of M. pacifica and M. okhotensis were highly variable and no temporal trend was detected. Comparison with field abundance data for Metridia spp. suggests that our estimates of egg production and hatchability are too low, despite care with experimental conditions. All the Neocalanus species utilize the bloom as energy for juvenile growth. Neocalanus cristatus developed from C2 through C4, and stage duration of C3 was estimated to be 24 days. Neocalanus flemingeri also developed from C1 through C3, and stage durations of C1 and C2 were estimated to be 7–9 days. Neocalanus plumchrus occurred in small numbers from mid-April onward. The stage duration estimates for Neocalanus spp. are similar to those reported from the high-nutrition southeastern Bering Sea shelf. Gut pigment variation clearly showed nocturnal feeding by Metridia spp., while no diel changes in gut pigment were recognized for E. bungii or Neocalanus spp. The diel changes in gut pigment of Metridia spp. were related to their diel vertical migrations. The calendar of sequential responses of copepods to the phytoplankton bloom is summarized.

Human forcing of the copepod–fish–jellyfish triangular trophic relationship

Copepods, the most abundant planktonic metazoans, constitute an intermediate trophic position between phytoplankton and higher trophic-level animals such as fish and jellyfish. Fish and jellyfish are adversaries because they often compete for prey copepods and also can be prey of each other. The classical food chain represented by phytoplankton–copepod–fish is the main process leading to efficient and sustainable production of fish as human food. At present, more than 75% of world fish stocks are fully or over exploited. On the other hand, jellyfish populations have increased world-wide, particularly in waters under significant human influences. Two such cases are seen in East Asian waters, where massive blooms of moon jellyfish (Aurelia aurita s.l.) and giant jellyfish (Nemopilema nomurai) have repeatedly occurred in recent decades, causing severe damage to local fisheries. In this article, I will review the pivotal role of copepods in marine ecosystems, particularly in the Inland Sea of Japan, where the annual fish catch per unit area is among the world’s highest. Then, I will describe an ongoing ecosystem shift from dominance by fish to dominance by jellyfish as a consequence of human forcing. Finally, I will propose to create “sato-umi”, a coastal sea with high productivity and biodiversity with wise human interaction, where copepod production would most efficiently transforms into food for humans.

Differences in plankton community structure along the Godthåbsfjord, from the Greenland Ice Sheet to offshore waters

This study describes differences in plankton community structure and in chemical and physical gradients between the offshore West Greenland Current system and inland regions close to the Greenland Ice Sheet during the post-bloom in Godthabsfjorden (64° N, 51° W). The offshore region had pronounced vertical mixing, with centric diatoms and Phaeocystis spp. dominating the phytoplankton, chlorophyll (chl) a (0.3 to 3.9 µg l -1 ) was evenly distributed and nutrients were depleted in the upper 50 m. Ciliates and heterotrophic dinoflagellates constituted equal parts of the protozooplankton biomass. Copepod biomass was dominated by Calanus spp. Primary production, copepod production and the vertical flux were high offshore. The water column was stratified in the fjord, causing chl a to be concentrated in a thin sub-surface layer. Nutrients were depleted above the pycnocline, and Thalassiosira spp. dominated the phytoplankton assemblage close to the ice sheet. Dinoflagellates dominated the protozooplankton biomass, whereas copepod biomass was low and was dominated by Pseudocalanus spp. and Metridia longa. Primary production was low in the outer part of the fjord but considerably higher in the inner parts of the fjord. Copepod production was exceeded by protozooplankton production in the fjord. The results of both physical/chemical factors and biological parameters suggest separation of offshore and fjord systems.

Seasonal changes in food quantity and quality of the common North Sea copepods Temora longicornis and Pseudocalanus elongatus: a bioassay approach

We evaluated the food quantity and quality over a seasonal cycle for the development and egg production of the common North Sea copepods Temora longicornis and Pseudocalanus elongatus, using a bioassay approach. Seston was sampled from December to October from a well- mixed water column of the Marsdiep (Dutch Wadden Sea) and fed to cultured copepods at a constant temperature of 15°C, thus excluding seasonal effects of temperature, body size, age, and maternal nutrition. Copepod response was evaluated by measuring egg production and juvenile development, while the seston quantity and quality were measured as the concentrations of chl a, specific phyto- plankton pigments, particulate organic carbon (POC), particulate organic nitrogen (PON), fatty acids, and sterols. The egg production of both copepods was low when feeding on seston collected in win- ter, but increased to peak values with the seston from the spring bloom in March-April. The juveniles of both species were able to complete their development only in spring experiments. A multiple regression analyses and comparison to a good-quality standard food of the same concentration sug- gested that, in an annual scale, the egg production and development of T. longicornis mainly depended on phytoplankton concentration, while the egg production and development of P. elonga- tus appeared also to benefit from detritus or heterotrophic food sources. The present study did not detect an influence of a specific food quality variable; however, an unexplained high juvenile mortal- ity in summer suggests that all factors are not understood yet.

Carbon transfer in a herbivore- and microbial loop-dominated pelagic food webs in the southern Barents Sea during spring and summer

MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 398:93-107 (2010) - DOI: https://doi.org/10.3354/meps08335 Carbon transfer in a herbivore- and microbial loop-dominated pelagic food webs in the southern Barents Sea during spring and summer Frederik De Laender*, Dick Van Oevelen, Karline Soetaert, Jack J. Middelburg NIOO-KNAW, Centre for Estuarine and Marine Ecology, Korringaweg 7, PO Box 140, 4401 Yerseke, The Netherlands *Email: frederik.delaender@ugent.be ABSTRACT: We compared carbon budgets between a herbivore-dominated and a microbial loop-dominated food web and examined the implications of food web structure for fish production. We used the southern Barents Sea as a case study and inverse modelling as an analysis method. In spring, when the system was dominated by the herbivorous web, the diet of protozoa consisted of similar amounts of bacteria and phytoplankton. Copepods showed no clear preference for protozoa. Cod Gadus morhua, a predatory fish preying on copepods and on copepod-feeding capelin Mallotus villosus in spring, moderately depended on the microbial loop in spring, as only 20 to 60% of its food passed through the microbial loop. In summer, when the food web was dominated by the microbial loop, protozoa ingested 4 times more bacteria than phytoplankton and protozoa formed 80 to 90% of the copepod diet. Because of this strong link between the microbial loop and copepods (the young cod’s main prey item) young cod (<3 yr) depended more on the microbial loop than on any other food web compartment, as >60% of its food passed through the microbial loop in summer. Adult cod (≤3 yr) relied far less on the microbial loop than young cod as it preyed on strictly herbivorous krill in summer. Food web efficiency for fish production was comparable between seasons (~5 × 10–4) and 2 times higher in summer (5 × 10–2) than in spring for copepod production. KEY WORDS: Food web · Microbial loop · Protozoa · Copepods · Gadus morhua Full text in pdf format Supplementary Material PreviousNextCite this article as: De Laender F, Van Oevelen D, Soetaert K, Middelburg JJ (2010) Carbon transfer in a herbivore- and microbial loop-dominated pelagic food webs in the southern Barents Sea during spring and summer. Mar Ecol Prog Ser 398:93-107. https://doi.org/10.3354/meps08335Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 398. Online publication date: January 05, 2010 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2010 Inter-Research.

Summer production of copepods in the Barents Sea

Summer production of copepods in the Barents Sea

Use of flubendazole as a therapeutic agent against rotifers (Brachionus plicatilis) in intensive cultures of the harpacticoid copepodTisbe holothuriae

Copepods are well known to be the optimal live feed for most species of marine fish larvae. Still copepods are rarely used in marine hatcheries worldwide. Lack of efficient production techniques are among the reasons for this. Consequently, Artemia and rotifers are utilized in commercial settings. One problem in intensive production of copepods is contamination with rotifers. Rotifers have higher growth rates than copepods and consequently will compete out the copepods when accidentally introduced to the copepod production systems. Once contamination has occurred, the only cure has been to shut down production and subsequently use a therapeutic agent to eliminate all zooplankton in the system before restart with a stock culture free of rotifers. We tested flubendazole as a mean of controlling rotifers (Brachionus plicatilis) in intensive laboratory cultures of the harpacticoid copepod (Tisbe holothuria). Flubendazole was lethal to rotifers in concentrations as low as 0.05 mg L−1. There was no significant effect on the concentration of copepods, even at the highest concentration tested, i.e. 5.0 mg L−1 flubendazole. We conclude that flubendazole is an effective drug for control of B. plicatilis in T. holothuriae batch cultures.

Variation in diatom biochemical composition during a simulated bloom and its effect on copepod production

The biochemical quality of phytoplankton depends in part upon nutrient availability, which has implications for the population dynamics of grazers. Here, we examined how nutrient availability influenced the growth dynamics and biochemical content of the marine diatom Thalassiosira weissflogii and how these biochemical changes impacted the vital rates of a calanoid copepod (Acartia tonsa). Changes in biochemistry protein, carbohydrate and fatty acids) were compared in diatom cultures that simulated bloom conditions (B-algae) and those maintained in near exponential growth (E-algae) over the course of a 16-day experiment. Egg production rates (EPRs, eggs female -1 day -1 ) and the developmental success of copepodite stages of A. tonsa fed these different diets were quantified. Copepod EPR was significantly lower (reduced by half) when B-algae entered the senescent phase due to silicate limitation. In a crossover (diet switch) experiment, EPR increased when copepods fed B-algae were switched to E-algae and vice versa. Copepodites of A. tonsa developed normally and reached the adult (C6) stage when fed E-algae, but ceased development (approximately at stage C2) when reared on senescent phase B-algae. Given the importance of copepods as prey for higher trophic levels, our results highlight how nutritional changes that naturally occur during a phytoplankton bloom may influence the productivity of copepods and higher trophic levels.

Copepod biomass, size composition and production in the Southern Benguela: Spatio–temporal patterns of variation, and comparison with other eastern boundary upwelling systems

Zooplankton sampling has been conducted biannually off the west and south-west coasts of South Africa since 1988, as part of the pelagic fish stock assessment cruises covering the continental shelf (0–200 m). Previously established relationships between body mass and growth rate were used to derive copepod production. Mean copepod biomass and annual production in the Southern Benguela between 1988 and 2003 were 2.1 g C m −2 and 134.4 g C m −2 yr −1, respectively. Biomass and production were higher during summer in four latitudinal areas, but seasonal variability was only significant for the central West Coast. This area, lying downstream from the Cape Columbine upwelling cell, supported the highest levels of copepod biomass (3.1 g C m −2) and production (545 g C m −2 d −1). Production during winter, when upwelling is minimal, contributed over one third of annual production in all four areas. Copepod biomass, daily production and size composition varied with latitude, season and year, and there were no consistent trends in interannual variability over the study period. Although biomass in the four major EBUS was of the same magnitude (usually <5 g C m −2), annual production estimates are an order of magnitude greater than those made four decades ago, with highest values for the Southern Benguela.

Feeding ecology of myctophid fishes in the northern Scotia Sea

The diets of 9 species of myctophid fishes, Electrona carlsbergi, E. antarctica, Gymno- scopelus fraseri, G. nicholsi, G. braueri, Protomyctophum bolini, P. choriodon, Krefftichthys ander- ssoni and Nannobrachium achirus, were investigated during austral autumn in the northern Scotia Sea. Based on the percent index of relative importance (%IRI), the data suggest dietary specialisation in some species, which may permit resource partitioning. Hierarchical agglomerative cluster analy- sis of Bray-Curtis similarity (60% threshold) separated the myctophid community into distinct feed- ing guilds. One group (G. braueri and E. antarctica) fed principally on Themisto gaudichaudii, another (P. choriodon and G. fraseri) primarily on copepods (Metridia spp. and Rhincalanus gigas), and a third group (G. nicholsi and P. bolini) consumed copepods and euphausids (mostly Metridia spp. and Euphausia frigida). The diets of E. carlsbergi and K. anderssoni differed from the other spe- cies, with E. carlsbergi being the only species that consumed salps. There was a general switch in diet from copepods to euphausiids and amphipods as the myctophid predator size increased. Dietary spe- cialisation is likely the result of a combination of predator size, gape size, filtering capacity of the gill rakers and the vertical distribution of predators and prey. Antarctic krill were only consumed by the larger myctophids, which represented a numerically minor part of the myctophid community, sup- porting the concept that myctophids can provide a krill-independent link between secondary produc- tion and higher trophic levels. However, the northern Scotia Sea is dominated by adult krill, which are only suitable as prey for larger fish. In the northern Scotia Sea, myctophid predation had a very small impact on copepod production but a higher impact on macrozooplankton, with a best-estimate of 4% of the daily production of Themisto gaudichaudii and 6% of that of Euphausia superba being consumed.

Estuarine and oceanic influences on copepod abundance and production of a subtropical coastal area

We investigated the influence of nutrient-rich oceanic waters in comparison to the estuarine outflow from Santos Bay (SE Brazil) on copepod abundance and production on the adjacent inner shelf. Zooplankton samples were collected with a Multinet in spring 2005 and in summer 2006. Copepod biomass was derived from length―weight regressions, and growth rates were estimated from empirical models. Altogether, 58 copepod taxa were identified. The highest abundances were due to small-sized organisms including nauplii, oncaeids and copepodids of paracalanids and clausocalanids. Biomass and secondary production mirrored copepod abundance, with Temora copepodids accompanying the above-mentioned taxa as major contributors. The contribution of naupliar biomass and production was low (2.2 and 3.8% of the total, respectively). The influence of the Santos Bay outflow was observed only in spring when Coastal Water (CW) dominated at the study site; whereas in summer the inner shelf was occupied by CW in the suface layer and the oceanic South Atlantic Central Water (SACW) in the bottom layer. The SACW intrusion had more of an influence for the increase in copepod production than the Santos Bay plume. The distribution and dynamics of the oceanic water masses seemed to be the most important influence on copepod diversity and production at this subtropical site.

An individual-based model to study the reproduction of egg bearing copepods: Application to Eurytemora affinis (Copepoda Calanoida) from the Seine estuary, France

Limited empirical studies have elucidated the daily egg production and associated reproductive processes of egg bearing copepod. Herein, we present an individual-based model which constitutes a realistic representation of the reproduction in egg bearing copepods. The model has been parameterized using an extensive set of experimental data obtained from the literature and from the laboratory and field experiments on the estuarine copepod Eurytemora affinis. The proposed model takes into account the adult female longevity, the clutch size and interclutch duration, which is a function of egg maturation time and latency time required by the female after egg hatching to produce a new clutch. The embryonic development time and hatching success are also taken into account. The effect of temperature on the means and variances of above-mentioned reproductive parameters has been also incorporated. A multi agent system based generic platform “ Mobidyc” has been used to generate and calibrate the model. The model demonstrates the reproductive parameters of females of E. affinis which is validated through individual based experiments. Temperature specific simulations provide a dynamical explanation of temperature effect on the cumulative egg production. The daily survival principally affects the number of clutches produced per female during its life span. The results obtained in the present study by combining temperature and survival effects reveal the relatively greater importance of the first factor on the daily egg production of egg-carrying copepods. The present model is generic and hence easily applicable to other animals with comparable reproductive strategy.

Aldehyde-induced insidious effects cannot be considered as a diatom defence mechanism against copepods

The last 2 decades have seen much controversy concerning the negative impacts that consumption of diatoms may have upon copepods. Because diatoms have traditionally been consid- ered a major food item for copepods, this controversy has raised fundamental questions concerning our understanding of the marine food chain. The mode of operation of this negative impact is via so- called insidious effects on the survival of early-stage copepod progeny, caused for example by polyunsaturated aldehydes (PUA), with the mechanism proposed as a defence evolved by diatoms. Here the effects of diatoms on copepod reproduction rates are evaluated using a stoichiometric weight- or age-stage copepod model coupled with variable stoichiometric descriptions of prey. We show that, considering different combinations of copepod population age structure and diet compo- sition (diatoms, non-diatoms, microzooplankton and copepods), the concept of an 'insidious' effect of diatom consumption upon copepods cannot be sustained as a defence mechanism, as it does not con- fer any advantage to the diatoms. While metabolites that deter grazing can be advantageous, slowly killing a predator (via the death of juvenile copepods) responsible for consuming competitor primary producers, and also consuming another predator (microzooplankton), is not advantageous to the diatoms. Indeed it is shown to be counterproductive on occasion. While PUA (or compounds that act in a similar fashion) have the potential to adversely affect copepod production, alternative explana- tions for their production need consideration. Such alternatives include the potential impact of PUA against microzooplankton, or simply that PUA represent secondary metabolites of no evolutionary selective advantage.

Limitation of egg production in Calanus finmarchicus in the field: A stoichiometric analysis

The egg production of marine copepods correlates with a range of variables, including the availability of organic carbon (C), nitrogen (N) and the polyunsaturated fatty acids (PUFAs) 20:5( n−3) (EPA) and 22:6( n−3) (DHA). However, an understanding of which substrates limit egg production in the natural environment has yet to be reached. The quantities of C, N, EPA and DHA ingested, derived from parental biomass, and invested in eggs by female Calanus finmarchicus during a 5-day incubation experiment were examined using stoichiometric theory to determine which substrate was limiting. The majority of each substrate was derived from parental biomass, and therefore the existing stoichiometric theory is developed to include this route of supply. The females were essentially devoid of lipid reserves, as evidenced by the lack of the storage fatty acids 20:1( n−9) and 22:1( n−11), and carbon limitation was predicted under most of the scenarios examined. Nitrogen limitation was only apparent when carbon and nitrogen utilisation efficiencies were assumed to be high (0.5) and low (0.4) respectively. PUFAs were assumed to be utilised with high efficiency (0.9), and were never predicted to limit production. This work highlights the need for a more detailed understanding of the maintenance requirements that marine copepods have for C, N, EPA, and DHA and hence the efficiencies with these substrates can be utilised for growth.

Metazooplankton distribution across the Southern Indian Ocean with emphasis on the role of Larvaceans

The abundance and depth distribution of metazoans >20 μm were investigated at seven stations across the Southern Indian Ocean (SIO), October–November 2006. Copepod nauplii, copepodites and larvaceans dominated the metazooplankton community. Copepodites were most abundant within Agulhas Current and Southern Ocean waters, decreasing toward subtropical/tropical areas, whereas larvaceans showed the inverse pattern. The fraction <200 μm contained the majority of the zooplankton enumerated, including 81, 23 and 93% of the larvacean, copepodite and nauplii abundances, respectively. The relative abundance of larvaceans compared with copepodites increased from 7 to 44% from South Africa towards Australia. Peak copepodite biomass was observed off South Africa, while larvacean biomass was <1% of the copepodite biomass there, increasing to 6% in tropical waters. Both copepodite and nauplii biomass were positively correlated to total Chl a (P < 0.0001), larvacean biomass was only significantly related to temperature (P = 0.0213). Despite their low biomass, larvacean production was estimated to exceed the copepod production up to five times. It appears that the abundance and role of larvaceans in the SIO has been severely underestimated in previous studies; thus future investigations into the fate of organic matter will remain incomplete if this group is not adequately considered.