Spatio-temporal distribution and production of calanoid copepods in the central Baltic Sea

The aim of our study was the exploration of species-specificdistribution and production patterns of dominant copepods inthe Central Baltic Sea (Bornholm Basin). Spatio-temporal distribution,egg and secondary production were studied by means of net-samplingand egg production experiments from April to August 1999. Verticaland horizontal distribution patterns appeared to be linked witheach other and were primarily species-dependent. Both Acartiaspp. and Temora longicornis preferentially inhabited the upper30 m of the water column and the shallower marginal regionsof the Bornholm Basin. In contrast, the C4–5 and C6 ofPseudocalanus spp. preferentially inhabited the halocline region(50–70 m) and the deep central part of the area. Observeddifferences in horizontal distribution among these three copepodsappear to result from different depth preferences and depth-dependentwater circulation patterns. Egg production rates (EPR) clearlyexhibited seasonal differences with maximum EPR in May and minimumEPR in July. Mean EPR was significantly different between species,being highest in Acartia spp. and lowest in Pseudocalanus spp.The variability in EPR was related to differences in hydrographyand modelled food environment.     

Capelin larvae (Mallotus villosus) and community structure of zooplankton off the coast of northern Norway

Capelin larvae (Mallotus villosus) and mesozooplankton werecollected in May 2001 in the southern part of the Barents Sea.During the period of hatching and first feeding, capelin larvaeexperience high mortality rates. In order to understand theunderlying mechanisms influencing larval survival, we monitoreda selected site for 10 days using a variety of different samplingapproaches. Hydrological parameters conductivity–temperature–depth–fluorescence(CTDF) were recorded and zooplankton samples obtained simultaneouslyusing MOCNESS at three time periods. The mesozooplankton consistedessentially of copepods (85), dominated by Calanus finmarchicus.The zooplankton community consisted of different assemblagesderived from different areas (i.e. different water masses; Atlanticand coastal). Capelin larvae were found in surficial coastalwaters. Copepod nauplii and C. finmarchicus were less abundantin coastal waters where small copepodites (such as Acartia spp.and Temora longicornis) were found together with early stagesof coastal invertebrates that are possible prey for capelinlarvae. We suggest that capelin larvae base their foraging onsmall prey organisms frequently found among the coastal zooplanktonand that these larvae are less dependent on the propagationof the recruiting generation of C. finmarchicus present fromApril to June.     

Zooplankton response to a warmer northern Wadden Sea

Weekly measurements of mesozooplankton (>76 μm) and hydrographic parameters have been carried out since 1984 in the List Tidal Basin (northern Wadden Sea). Monthly water temperature significantly increased by 0.04°C year⁻¹. The largest increase by 3°C in 22 years occurred in September, implying an extension of the warm summer period. Mean annual copepod abundance and length of copepod season correlated significantly with mean temperature from January to May. Except for an increasing Acartia sp. abundance during spring (April–May), no long-term trends in copepod abundance were observed. The percentage of carnivorous zooplankton increased significantly since 1984 mainly due to a sudden increase in the cyclopoid copepod Oithona similis in 1997. We expect that global warming will lead to a longer copepod season and higher copepod abundances in the northern Wadden Sea.     

Mesozooplankton structure in Dolgaya Bay (Barents Sea)

The community structure of mesozooplankton was investigated in Dolgaya Bay (southern Barents Sea), a subarctic fjord, using a Juday net (0.168-mm mesh size) in July, 2008. A total of 39 species and higher taxa were found. Average abundance, biomass and diversity (±standard error) were 153,403 ± 15,855 ind m⁻², 570 ± 61 mg dry mass m⁻², and 2.25 ± 0.09, respectively. Copepods were the most numerous species. The mesozooplankton communities were dominated by omnivores (Oithona similis and Acartia spp.). Vertical distribution of the mesozooplankton was characterized by copepod dominance at each sampled layer. There were no significant correlations among physical variables and biological parameters, except negative correlation for the mean biomass and mean water temperature.     

Zooplankton diversity and the predatory impact by larval and small juvenile fish at the Fisher Banks in the North Sea

The biomass and diversity of the mesozooplankton and fish larvaecommunity were investigated across a frontal zone in the centralNorth Sea in the early summer, to investigate whether larvalfish predation is a regulator of mesozooplankton production.Pronounced changes in the mesozooplankton community were observedacross the front off the Jutlandic coast. The biomass and thediversity of copepods changed across the front as the populationof Calanus finmarchicus became abundant in the deeper water.The crustaceans (Acartia spp. and Evadne spp.) and polychaetelarvae dominated the mesozooplankton in the coastal water. Thebiomass of fish larvae was dominated by gadoid larvae. As inthe copepods, a shift in fish diversity was observed in thefrontal zone. On the coastal side of the front, whiting (Merlangiusmerlangus) dominated the biomass, while offshore from the frontwhiting were absent and cod (Gadus morhua) was the dominantlarval fish species on the deeper stations. The present investigationdemonstrates two different trophic pathways related to hydrographyin the central North Sea. First, in the shallow coastal water,the abundant small neritic copepods are predominately predatedby whiting, while in the offshore region the larger oceaniccopepods are predated by cod larvae. However, the predationpressure by the fish larvae was in general low (<10%) relativeto copepod production per day. Consequently, in the early summer,the copepod production potentially results in a build-up ofcopepod biomass on both sides of the front.     

Long-term dynamics of zooplankton in the southeastern Baltic Sea

Long-term research in the Baltic Sea revealed the basic trends of zooplankton community variations depending on oceanographic processes. Alternation of the periods of increase and decrease in salinity of the Baltic Sea against the background of climate changes (temperature increase) and eutrophication affect the state of the entire Baltic ecosystem, including zooplankton. For these periods, the dynamics of zooplankton in the Baltic Sea were analyzed based on literature data and results of regular research in the southeastern Baltic Sea during 1998–2007. The changes in the hydrological situation were accompanied by significant changes in the zooplankton community. In the 1990s–2000s, the abundance and biomass of brackish-water and thermophilous species primarily of Cladocera and Copepoda increased markedly. The role of the previously dominant marine copepod Pseudocalanus elongatus decreased due to salinity reduction in the deep-water part of the Baltic Sea. Maximum development of zooplankton occurred in years of the greatest warming-up of the water (2001, 2005–2007) against the background of a general positive trend of zooplankton abundance in the last decade.     

Copepod grazing in a summer cyanobacteria bloom in the Gulf of Finland

Blooms of the cyanobacteria Nodularia spumigena and Aphanizomenon flos-aquae dominated the phytoplankton assemblages of the western Gulf of Finland and the eastern side of the northern Baltic Sea in late July–August, 1992. The bloom overlapped the peak seasonal contributions of the dominant mesozooplankton herbivores in the region, the copepods Acartia bifilosa and Eurytemora affinis and the cladoceran Bosmina longispina maritima. Using radio-labelling techniques; the copepods were offered one of the cyanobacteria, Nodularia, as well as the 10–54 µm fraction of the natural phytoplankton assemblage. In general, incorporation rates of the labelled phytoplankton into the copepods declined with increasing contributions of the cyanobacteria. For both copepods, incorporation was inversely related to total phytoplankton biomass, whether measured as chlorophyll, total cells or cyanobacteria biomass. The very low rates for Acartia (< 0.8 µl [copepod h]^–1) indicated that this copepod was likely starving in the cyanobacteria bloom, consistent with the generally poor condition of the animal observed in the laboratory. The other major mesozooplanktor, B. longispina maritima, ingested substantially more cyanobacterial biomass than the two copepods, based on HPLC-identified cyanobacteria-specific pigment echinenone in the gut. Bloom carbon provided < 1% and < 4% of the daily rations for Acartia and Eurytemora, respectively. Total copepod demand in the cyanobacteria blooms was trivial, < 1% of bloom biomass consumed daily. These results suggest that copepod herbivory is relatively unimportant in dissipating summer cyanobacteria blooms in the Gulf of Finland.     

Carbon, nitrogen and phosphorus stoichiometry of crustacean zooplankton in the Baltic Sea: implications for nutrient recycling

The carbon (C), nitrogen (N) and phosphorus (P) contents (%of dry weight) of some crustacean zooplankton were studied inthe Baltic Sea. The copepod Acartia sp. had a stable C and Ncontent (48.3 ± 0.8% C, 12.4 ± 0.2% N, C:N ratio4.5 ± 0.1). The P content was variable (1–2%),probably depending on developmental stage and season. Copepodsaccumulating fat, like Pseudocalanus minutus elongatus, hadhigher and more variable C content (50–60%), and lowerN and P content (7–12% N, 0.6–1.5% P). The highestC and lowest N and P contents were found in adult Limnocalanusmacrurus. However, the N:P ratio was apparently independentof fat content and between 14 and 27 for all copepods. The cladoceransBosmina longispina maritima and Evadne nordmanni had lower Ncontent (9.3–10.8%) and higher C:N ratio (5.1–5.7)than Acartia sp. The P content (1.2–1.4%) was similarto Acartia sp. and the N:P ratios (16–19) were in thelower range of that found for the copepods. The N:P ratio wasgenerally somewhat higher in the copepods than in seston, whichmost of the year had nearly Redfield C:N:P ratios. Potentially,nutrient recycling from crustacean zooplankton could enhanceN limitation of phytoplankton, but small stoichiometric differencessuggest that this effect is probably weak. The extent is dependenton the structure of the zooplankton community and the grossgrowth efficiencies. Acartia copepodites, which had nearly RedfieldN:P ratios, would have the opposite effect and enhance P limitationin late summer when seston N:P ratios increased.     

Predation impact of age-0 fish on a copepod population in a Baltic Sea inlet as estimated by two bioenergetics models

Decreasing spring abundances of the copepod Eurytemora affinis,in a shallow brackish inlet of the Baltic Sea, were comparedwith calculated consumption of age-0 fish [dominant speciesherring (Clupea harengus) and perch (Perca fluviatihs)] as determinedby two bioenergetics models. Fish and zooplankton populationswere sampled at three stations along a salinity gradient of0.3-8.1% maximum range. Larvae and juveniles of both fish speciespreferred to prey on adults and developmental stages of E.affinis.Although there were substantial variations in fish density andcopepod production across the salinity range, consumption ofzooplankton by age-0 fish was never high enough to be a majorsource of copepod mortality.This finding is an independent confirmationof previous results obtained by direct food-intake measurementsin the same region. The low impact of fish predation may bea function of the low individual fish biomass in spring, theincreased copepod production and the dominance of the less planktivorousperch, rather than herring, in the less saline waters of theBarther Strom inlet.     

Metabolism and foraging strategies of mid‐latitude mesozooplankton during cyanobacterial blooms as revealed by fatty acids,amino acids,and their stable carbon isotopes

Increasing sea surface temperatures (SST) and blooms of lipid‐poor, filamentous cyanobacteria can change mesozooplankton metabolism and foraging strategies in marine systems. Lipid shortage and imbalanced diet may challenge the build‐up of energy pools of lipids and proteins, and access to essential fatty acids (FAs) and amino acids (AAs) by copepods. The impact of cyanobacterial blooms on individual energy pools was assessed for key species temperate Temora longicornis and boreal Pseudo‐/Paracalanus spp. that dominated field mesozooplankton communities isolated by seasonal stratification in the central Baltic Sea during the hot and the cold summer. We looked at (a) total lipid and protein levels, (b) FA trophic markers and AA composition, and (c) compound‐specific stable carbon isotopes (δ¹³C) in bulk mesozooplankton and in a subset of parameters in particulate organic matter. Despite lipid‐poor cyanobacterial blooms, the key species were largely able to cover both energy pools, yet a tendency of lipid reduction was observed in surface animals. Omni‐ and carnivory feeding modes, FA trophic makers, and δ¹³C patterns in essential compounds emphasized that cyanobacterial FAs and AAs have been incorporated into mesozooplankton mainly via feeding on mixo‐ and heterotrophic (dino‐) flagellates and detrital complexes during summer. Foraging for essential highly unsaturated FAs from (dino‐) flagellates may have caused night migration of Pseudo‐/Paracalanus spp. from the deep subhalocline waters into the upper waters. Only in the hot summer (SST>19.0°C) was T. longicornis submerged in the colder subthermocline water (~4°C). Thus, the continuous warming trend and simultaneous feeding can eventually lead to competition on the preferred diet by key copepod species below the thermocline in stratified systems. A comparison of δ¹³C patterns of essential AAs in surface mesozooplankton across sub‐basins of low and high cyanobacterial biomasses revealed the potential of δ¹³C‐AA isoscapes for studies of commercial fish feeding trails across the Baltic Sea food webs.     

The influence of individual variation on length-biomass regressions in three crustacean zooplankton species

Length-biomass regressions were established for the cladoceranBosmina longispina maritima and for the copepods Euryremoraaffinis hirundoides and Limnocalanus macrurus, all common inthe Baltic Sea. Biomass was analysed as individual organic carboncontent. For Bosmina, length was a weak estimator of biomass;only 51% of the variation in carbon content was explained bylength. For pre-adult stages of copepods length was a rathergood estimator of biomass. For Euryremora nauplii. the length-carbonrelationship was best described by linear equations (r² wasbetween 0.49 and 0.69), and for copepodite stages (CI-V) byexponential functions (r² = 0.81). The regressions for two differentsampling areas (Baltic proper and Bothnian Bay) did not showstatistically significant differences. Among the copepodites(CI-V) of Limnocalanus there was a linear relationship betweenlength and carbon content (r² = 0.76). In both copepod speciesthere was no or a weak correlation between length and biomassamong adults.     

Carbon content of some common tropical Andaman Sea copepods

Individual carbon contents of eight common copepod speciesfrom the Andaman Sea were determined. Length–carbon regressionsare presented for four calanoids (Acrocalanus gibber, Centropagesfurcatus, Temora discaudata and Euchaeta spp.), two cyclopoids(Oncaea spp. and Corycaeus spp.) and two harpacticoids (Macrosetellagracilis and Microsetella spp.). The copepod specimens wereobtained from different localities and times of the year during1996–1997. The regression coefficients are good in calanoidand cyclopoid copepods, but poor in the harpacticoids. The slopevalues range from 2.3 to 3.8 in the calanoids to 1.2 and 1.6in the harpacticoids, while the cyclopoids have slope valuesof 2.0 and 2.9. The equations derived from this study are expectedto be general characteristics, which are applicable for thecalculation of copepod biomass and production in tropical areas.     

Copepod feeding behaviour and egg production during a dinoflagellate bloom in the North Sea

Feeding strategies of copepods were studied during a dinoflagellate-dominated bloom in the North Sea in August 2001. The aim of this study was to evaluate the importance of mesozooplankton grazing as a biological loss factor of harmful algal blooms under natural conditions. Therefore, ingestion, egestion and egg production experiments were performed with the most abundant copepod species Calanus helgolandicus, Temora longicornis and Acartia sp. feeding on the natural phytoplankton community. Dinophysis norvegica and Ceratium furca were the most abundant dinoflagellate species at the time of the experiments. Grazing experiments as well as examination of fecal pellet content revealed C. helgolandicus fed efficiently on D. norvegica. Ingestion rates up to 47 cells female⁻¹ h⁻¹ were measured and a large proportion of the C. helgolandicus fecal pellets contained intact D. norvegica cells. Dinophysis cells were rarely seen in fecal pellets produced by T. longicornis, and never observed in pellets produced by Acartia sp. The ingestion rate of C. furca, which was the dominating Ceratium species, mimicked that of D. norvegica. C. helgolandicus grazed significantly on C. furca (16 cells female⁻¹ h⁻¹), while the ingestion rate of T. longicornis was low and Acartia sp. was not able to graze on C. furca. Egg production experiments revealed that 92% of the C. helgolandicus females produced eggs. The specific egg production rate and the proportion of females producing eggs among T. longicornis were low. This field experiment clearly shows that some copepod species feed efficiently on D. norvegica and C. furca under natural conditions, which may affect the bloom development of these dinoflagellates.     

东海中华哲水蚤的年产量估算

1 997年至 2 0 0 0年在东海陆架区进行了 4个航次的大面调查 ,用获得的大型浮游生物网样品分析资料应用产量 /生物量比值 ( P/ B)法对中华哲水蚤的年产量做了初步估算。中华哲水蚤的 P/ B比值为6.735 3,在东海的平均年产量为 2 63.94C mg/ ( m2 / a)。全测区年产量为 1 1 0 .745× 1 0 3t碳 ,即 1 1万 t,相当于活体重 1 44 .3万 t。     

A Less Saline Baltic Sea Promotes Cyanobacterial Growth,Hampers Intracellular Microcystin Production,and Leads to Strain-Specific Differences in Allelopathy

Salinity is one of the main factors that explain the distribution of species in the Baltic Sea. Increased precipitation and consequent increase in freshwater inflow is predicted to decrease salinity in some areas of the Baltic Sea. Clearly such changes may have profound effects on the organisms living there. Here we investigate the response of the commonly occurring cyanobacterium Dolichospermum spp. to three salinities, 0, 3 and 6. For the three strains tested we recorded growth, intracellular toxicity (microcystin) and allelopathic properties. We show that Dolichospermum can grow in all the three salinities tested with highest growth rates in the lowest salinity. All strains showed allelopathic potential and it differed significantly between strains and salinities, but was highest in the intermediate salinity and lowest in freshwater. Intracellular toxin concentration was highest in salinity 6. In addition, based on monitoring data from the northern Baltic Proper and the Gulf of Finland, we show that salinity has decreased, while Dolichospermum spp. biomass has increased between 1979 and 2013. Thus, based on our experimental findings it is evident that salinity plays a large role in Dolichospermum growth, allelopathic properties and toxicity. In combination with our long-term data analyses, we conclude that decreasing salinity is likely to result in a more favourable environment for Dolichospermum spp. in some areas of the Baltic Sea.     

Mesozooplankton in the northern Wadden Sea of Sylt: Seasonal distribution and environmental parameters

A statistical analysis was performed to determine environmental factors of significant influence on the mesozooplankton in the northern Wadden Sea of Sylt (German Bight) in 1991. The total mesozooplankton (>76 μm) was significantly correlated with water temperature as was the case with the calanoid copepodAcartia sp.;Pseudo- andParacalanus sp. were influenced by the amount of rainfall, whereas the amount of the meroplanktonic larvae of the cirripedia was related to the amount of particulate organic nitrogen in the water. A comparison of the years 1984 and 1991 demonstrates the positive effect of temperature on the copepod speciesAcartia sp. in summer.     

Mesozooplankton distribution in relation to hydrology of the Northeastern Aegean Sea, Eastern Mediterranean

The Northeastern Aegean Sea (NEA) is the area where Black Seawaters (BSW) outflow in the Mediterranean enhancing local productivityand inducing high hydrographic complexity. We describe the structureand distribution (both vertical and horizontal) of mesozooplanktonassemblages in the NEA during the thermal stratification periodin July and September 2003, in an effort to identify BSW effects.The overall standing stocks of mesozooplankton in the NEA weremuch higher than those typically reported for other pelagicwaters of the eastern Mediterranean. Higher abundance and biomassvalues and a distinctive copepod and cladoceran species assemblagewere recorded in the surface layer (directly influenced by theBSW). Variability in the supply and subsequent advection ofBSW in the NEA seemed to be the major factor affecting the structureand distribution of mesozooplankton assemblages. In July, lowersurface salinity reflected the increased supply of BSW (richin organic carbon) and, consequently, the abundance of filterfeeding taxa (cladocerans, appendicularians and doliolids) washigher than in September. During both surveys, the copepod/cladoceranassemblages of the surface (0–50 m) layer were sensitivetracers of horizontal hydrological and biological variability(e.g. location of fronts) associated with the advection of theBSW. This paper was presented at Plankton Symposium III, held atFiguera da Foz, Portugal between 17 and 20 March 2005, underthe auspices of the University of Coimbra and the Universityof Aveiro, and coordinated by Mário Jorge Pereira andUlisses M. Azeiteiro.     

Exoskeletal scars caused by bacterial attachment to copepods

Indications of exoskeletal scars on copepods Acartia spp. suggestbacterial attachment. Infestation of > 10⁵ bacterial cellsper copepod may affect differences in respiratory rate for theanimals.     

Seasonal variability in copepod ingestion and egg production on the Faroe shelf

Copepod community ingestion rates of Calanus finmarchicus, Temora longicornis, Acartia longiremis and Pseudocalanus spp., and egg production rates of C. finmarchicus and T. longicornis, were studied in relation to phytoplankton composition, abundance and biomass on the Faroe shelf during a one-year cycle. The phytoplankton community during winter was mainly composed of small flagellates and the copepods of Pseudocalanus spp. As the spring bloom progressed, diatoms increased in abundance and dominated the biomass throughout summer. C. finmarchicus increased in numbers in early spring, while T. longicornis and A. longiremis dominated the community during summer and autumn. While no response in ingestion rates was observed for A. longiremis and Pseudocalanus spp. with increasing diatom biomass, both ingestion rates and egg production of C. finmarchicus and T. longicornis, generally increased, showing a dependence upon diatoms for production. The daily ingestion for C. finmarchicus females was 7% and 22% of body biomass during the pre-bloom and bloom period, respectively, while for T. longicornis females it was 33% and 56% and for A. longiremis females 22% and 33%, respectively. C. finmarchicus accounted for more than 80% of the total copepod ingestion in May, but in mid- and late summer, T. longicornis and A. longiremis dominated, and represented 80–90% of the total copepod ingestion. The proportion of reproductively mature C. finmarchicus increased as the phytoplankton biomass increased. Most of the time there was good agreement between herbivorous ingestion rates and calculated carbon demand for the observed egg production. However, both species showed a peak in egg production prior to the phytoplankton spring-bloom. Handling editor: J. Padisak.     

Effects of Food Availability and Predation on a Marine Zooplankton Community — a Study on Copepods in the Baltic Sea

The importance of food resources versus predatory effects in determining the abundance pattern of zooplankton varies over time and space. Here, we evaluate the population dynamics of dominant calanoid copepods (Acartia spp. bifilosa/A. longiremis and Eurytemora affinis) in a reference area and a eutrophied area in the northern Baltic Sea Proper. We base the paper on a combination of statistical analyses of a long term (12 years) data set on population biomass dynamics, laboratory experiments on the feeding biology of these species and literature data of their susceptibility to predation. Overall, copepods were equally abundant at both sites, but Eurytemora was more abundant in the eutrophied area while Acartia spp. dominated in the non-eutrophied reference area. There was, however, no significant difference between the two areas in biomass dynamics (biomass specific rate of change) of the two groups of copepods. However, in spring and early summer the population biomass increased faster for Eurytemora as compared to Acartia spp. This is consistent with our findings from feeding and respiration experiments. Eurytemora exhibited generally higher ingestion rates and had a higher growth efficiency (ratio of ingestion to respiration rates), which suggests an advantage over Acartia spp. when food availability increases. Both species generally showed a rapid population decline starting in late summer, most likely caused by intensive predation by fish and mysid shrimps. In late autumn and winter, biomasses of both genera continued to decrease, but the decline was fastest for Eurytemora. During this period of the year, when the food availability is sparse, Acartia spp. may have an advantage over Eurytemora, as we found that the former ingests a broader size range of food particles and it has also been reported to be able to switch from filter feeding to raptorial feeding. This study shows that dominant copepods in the net-zooplankton community in our study area were structured both by food availability and predation. The relative importance of these effects varied seasonally.