Population dynamics of calanoid copepods and the implications of their predation by clupeid fish in the Central Baltic Sea

Population dynamics of major Baltic calanoid copepod speciesin the Gotland Basin during the last two decades were characterizedby a decline of Pseudocalanus elongatus associated with decliningsalinities, and an increase of Temora longicornis and Acartiaspp. potentially due to warmer conditions. Additionally thisstudy investigated the effect of predation by the major planktivorousfish species herring (Clupea harengus) and sprat (Sprattus sprattus)for the period 1977–1996 in the Gotland Basin (CentralBaltic Sea). Examination of consumption by these fish speciesin relation to copepod production estimates showed a switchby herring from consuming mainly CV/VI of P. elongatus and T.longicornis, to preying on CII of the latter copepod. This switchwas potentially due to increased competition with the drasticallyincreased sprat stock since the late 1980s. Further, an increasedpredation pressure by sprat on CV/CVI of both copepod speciesin spring resulted in higher copepod mortality rates. In consequence,based on these results we suggest that the increase in the spratstock since the late 1980s contributed to a decline of P. elongatus,and additionally prevented an even more pronounced temperature-drivenincrease in the T. longicornis stock, as was observed for Acartiaspp., which was not significantly consumed.     

The distribution of calanoid copepods in the plankton of Wisconsin Lakes

Zooplankton communities from 499 lakes were examined for calanoid copepod species. Limnocalanus macrurus and Senecella calaloides are confined to Lakes Michigan, Superior and one inland lake each. Eurytemora affinis has recently become established in the coastal waters of Lakes Michigan and Superior. Epischura lacustris is present in the summer plankton of a wide variety of lake types. Aglaodiaptomus leptopus is sensitive to fish predation and confined to small lakes without fish. Leptodiaptomus ashlandi is restricted to Lakes Michigan and Superior, but L. sicilis is also found in some inland lakes. Leptodiaptomus minutus is a boreal species found mostly in lakes of the far north. Skistodiaptomus oregonensis is the most commonly occurring calanoid and seems well adapted to lakes of the meso-to low eutrophic types. Skistodiaptomus pallidus is also generally distributed throughout the state, but less common than oregonensis and adapted to more eutrophic lakes. Leptodiaptomus siciloides and A. clavipes are western species, adapted to eutrophic lakes, which are expanding their ranges eastward, aided by cultural eutrophication.     

Spatio-temporal distribution of Oithona similis in the Bornholm Basin (Central Baltic Sea)

Oithona similis is an abundant but poorly studied cyclopoidcopepod in the brackish Central Baltic Sea. We describe thespatio-temporal distribution of O. similis in a Central Balticdeep basin (Bornholm Basin) during spring and summer 1999. Usingvertically resolving sampling in parallel with hydrographicmeasurements, we found the copepod to dwell in the permanenthalocline characteristic of a Central Baltic deep basin. Thehabitat of O. similis is thus limited from above by low salinityand from below by low oxygen conditions, both characteristicfor the area. Horizontally resolving sampling yielded abundancesurfaces which were compared by analysis of variance showingsimilar patterns among sampling dates. Comparison with flowfields from a three-dimensional hydrodynamic model suggeststhat the horizontal distribution is primarily the result ofcirculation in the dwelling depth. The study shows how the physicalenvironment in the area determines the spatial distributionwhich might affect abundance and production of this copepod.     

Spatio-temporal dynamics of chlorophyll in the open Baltic Sea

Five charts of the chlorophyll and hydrographic fields by verticalprofiles of in situ fluorescence and CTD were made on a stationarygrid of 20 ? 25 nautical miles with a 5-mile spacing in theopen Baltic Sea. Both chlorophyll levels and variability weremaximal close to the spring bloom. High chlorophyll levels insummer are sustained by recurrent nutrient injections from thedeep saline layer. Two of the surveys showed close couplingbetween the coarse-scale (10 km) chlorophyll distribution andthe hydrographic structure determining the intensity of nutrienttransfer. Vigorous advection, stirring and current shear, associatedwith a strong mesoscale eddy, probably dominated the chlorophyllpattern on three surveys. The upward velocities in the cycloniceddy resulted in accumulation of phytoplankton in the aphoticzone. Intensive heat input from the surface caused a suddensinking of the phytoplankton and the formation of a pronouncedsub-surface chlorophyll maximum.     

Observations on the taxonomic composition and vertical distribution of cyclopoid copepods in the central Red Sea

The taxonomic composition and vertical distribution of cyclopoid copepods, including very small species, in the central Red Sea were analyzed. Samples were taken in a multiple opening and closing net with 0.1 mm mesh size to a depth of 450 m. Most species belong to the genus Oncaea (13 identified and 6 unidentified forms). Nearly one half of these are smaller than 0.4 mm in length. Nine cyclopoid species and three groups of very similar species and forms of Oncaea predominate. In the upper 450 m of the water column, each of these species or groups account for more than 1 % of all cyclopoid specimens. In the epipelagic zone (0–100 m) Oithona simplex predominates, accounting for 20 % of the total. In the upper mesopelagic zone Paroithona sp. and one of the unidentified Oncaea forms are most numerous from 100 to 250 m, and two groups of Oncaea are most abundant from 250 to 450 m. The dominance of single species among cyclopoids is less pronounced than that reported for calanoids in the mesopelagic zone of the central Red Sea.     

Lipid dynamics and feeding of dominant Antarctic calanoid copepods in the eastern Weddell Sea in December

Lipid content, fatty acid composition, and feeding activity of the dominant Antarctic copepods, Calanoides acutus, Calanus propinquus, and Metridia gerlachei, were studied at a quasi-permanent station in the eastern Weddell Sea in December 2003. During 3 weeks of the spring phytoplankton development, total lipid levels of females and copepodite stages V (CVs) of C. acutus were almost doubled. Meanwhile, only a slight increase in total lipid content occurred in M. gerlachei, and no clear trend was observed in lipids of C. propinquus females. The pronounced increase of lipids in C. acutus was due to an accumulation of wax esters. The proportion of wax esters in the lipids of M. gerlachei was clearly lower, while triacylglycerols played a more important role. In C. propinquus, triacylglycerols were the only neutral lipid class. There were no pronounced changes in the feeding activity of M. gerlachei, whereas the feeding activity of C. acutus had rapidly increased with the development of the phytoplankton bloom in December, which explains its rapid lipid accumulation. The combination of gut content and fatty acid trophic marker analyses showed that C. acutus was feeding predominantly on diatoms. The typical diatom fatty acid marker, 16:1(n-7), slightly decreased and the tracer for flagellates, 18:4(n-3), increased in females and CVs of C. acutus. This shift indicates the time, when the significance of flagellates started to increase. The three copepod species exhibited different patterns of lipid accumulation in relation to their trophic niches and different duration of their active phases. The investigations filled a crucial data gap in the seasonal lipid dynamics of dominant calanoid copepods in the Weddell Sea in December and support earlier hypotheses on their energetic adaptations and life cycle strategies.     

Dormant eggs of calanoid copepods in sea-bed sediments of the English Channel and southern North Sea

Nauplii of Centropages spp., Labidocera wollastoni and Temoralongicornis emerged from incubated samples of sea-bed sedimentsover a period of 6 months. Dormant eggs have not previouslybeen noted in T. longicornis.     

The calanoid copepods of the Nile system

At least 11 or 12 calanoid taxa occur in the Nile system. One species is typical of the river and of Lake Chad (Th. galebi); two are restricted to Lake Victoria (Th. galeboides, T. stuhlmanni), but both might be only subspecies to more widespread species. One is restricted to Lakes Edward and George (T. worthingtoni) but is a little known species. No calanoids are on record from Lakes Albert and Kyoga, while the species reported from Lake Turkana (T. banforanus) is out of range, and almost certainly represents an erroneous record. Among the remaining species, four are East-African, ranging from the southern tip of the continent to the Ethiopian plateau and the Nile valley (P. schultzei, Th. mixtus, T. kraepelini, T. cf orientalis), while two are Sahelian species that span Africa from east to west (M. mauretanicus, T. processifer et ssp.).Two new synonyms are introduced. One new subspecies (T. processifier friedae) is described from the Ethiopian plateau. It is suggested that Tropodiaptomus orientalis (Brady, 1886), the type species of its genus, should not be considered a nomen dubium, but should be redescribed on its type female(s) and on topotypical males from Sri Lanka.     

Egg production and hatching success in the calanoid copepods Calanus helgolandicus and Calanus finmarchicus in the North Sea from March to September 2001

Spatial and seasonal egg production rates (E_r) and egg hatchingsuccess in the copepods Calanus finmarchicus and Calanus helgolandicuswere measured in the North Sea from March to September. Foodavailability was monitored by chlorophyll and protist concentrationsand three size fractions of seston fatty acids. Seasonal andspatial distribution and production differed between the species.Calanus finmarchicus was found only offshore of the 50-m isobath,with decreasing E_r (37–28 eggs female^–1 day^–1)from March to July. Calanus helgolandicus had two abundancepeaks, in spring and autumn, with a low in May during whichtime the highest E_r were observed (38 eggs female^–1 day^–1).At other times, E_r in C. helgolandicus remained lower than inC. finmarchicus (     

The lipid biochemistry of calanoid copepods

Calanus species, particularly those in high latitudes, can accumulate large oil reserves consisting predominantly of wax esters. These wax esters consist predominantly of 16:0, 20:1 (n–9) and 22:1 (n–11) fatty alcohols, mainly formed de novo by the animals from non-lipid dietary precursors, esterified with various fatty acids that are often polyunsaturated fatty acids and largely of dietary, phytoplanktonic origin. Wax ester formation is maximal in copepodite stages IV and V. The lipids are elaborated not primarily for buoyancy regulation but as a source of metabolic energy during overwintering, particularly for reproduction. Large quantities of wax esters are utilised for gonadal development when stage V copepodites mature to females. Development of stage V copepodites to males is not accompanied by wax ester utilisation but males consume large amounts of these lipids in physical activity during reproduction. The role of wax esters in the life history of calanoids is illustrated with particular reference to a comparison of Calanus finmarchicus and Metridia longa in Balsfjord, northern Norway.     

Abundance,population structure and vertical distribution of dominant calanoid copepods on the eastern Weddell Sea shelf during a spring phytoplankton bloom

The main emphasis of this study was to analyse the short-term development of abundance, population structure and vertical distribution of the dominant calanoid copepods during a phytoplankton bloom in the coastal area of the eastern Weddell Sea in December 2003. Microcalanus pygmaeus was by far the most abundant calanoid species. Metridia gerlachei, Ctenocalanus citer, Calanoides acutus, Calanus propinquus and the ice-associated Stephos longipes were also present in considerable proportions. The observed changes in the population characteristics and parameters of these species are described in detail and discussed in the context of the spring phytoplankton bloom. A conspicuous event occurring during the final stage of the study was the development of a strong storm. While the results suggest that this storm did not have any considerable influence on the populations of all other investigated copepod species, it very likely caused pronounced changes in the S. longipes population present in the water column. Before the storm, S. longipes was found primarily in the upper 100 m of the water column, and its population was dominated by adults (mean proportion = 41%) and the copepodite stage I (mean proportion = 30%). After the storm, the abundance increased considerably, and the copepodite stage I contributed by far the largest proportion (53%) of the total population indicating that the early copepodite stages probably had been released from the sea ice into the under ice water layer due to ice break-up and ice melt processes caused by the storm.     

Colonization of the pelagic realm by calanoid copepods

The evolution of calanoid copepods probably extends back into the mid-Paleozoic. Environmental change from the Paleozoic through to the Tertiary is reviewed. Turbidity, water clarity, oxygen, pelagic primary production, and tectonically induced changes in the morphology of the oceans are probably all important drivers of calanoid evolution and their invasion of the pelagic realm. Current views of the phylogeny of the Calanoida are presented as well as a review of some recent work on metabolic potential, female genital system, and nervous system. It is hypothesized that ancestors of the Arietelloidea and Diaptomoidea invaded the water column in the Devonian at a similar time to the Ostracoda and that the ancestors of the Calanoidea–Clausocalanoidea, with their myelinated axons, arose in the Permian during the major deep ventilation of the ocean. Present day distributions of some Diaptomidae, Centropagidae, and Calanidae suggest that these families successfully came through the Jurassic/Cretaceous expansion of the oxygen minimum zone and the K-T boundary event. Some Arietelloidea and Clausocalanoidea became secondarily benthic and may have survived the K-T boundary event in this environment. It is postulated that some Clausocalanoidea reinvaded the water column (e.g. Clausocalanidae, Aetideidae, Phaennidae, Scolecitrichidae) in the Tertiary after finding refuge in deep, low oxygen water away from the sea surface. It is possible that these hypotheses may be testable using genetic information in the near future.     

Body size and sexual size dimorphism in calanoid copepods

Patterns of sexual size dimorphism and body size in calanoid copepods are examined. We hypothesize that favorable conditions for development will result in large body size and high sexual size dimorphism among populations of a given species and that differences in this allometric relationship among species is governed by the male's role in insemination. We confirm that there is a greater advantage to large female size, normally the larger sex, when compared to males, hence leading to selection for developmental patterns favoring high size dimorphism. Individuals from populations of four centropagid copepod species were measured; other sizes were obtained from published sources. In the four species we examined, the relationships between prosome length and both clutch size and the ability to produce multiple clutches with one insemination were determined. Results show a trend toward hyperallometry in all centropagid species examined: sexual size dimorphism increases with increasing size. Large females produce larger clutches and more additional clutches on one insemination. That hyperallometry is not observed in diaptomid copepods may result from the greater role the male plays in reproduction. Males are needed for each clutch produced, hence the selective pressure to be larger is greater than that in the centropagidae.     

Sexual dimorphism in calanoid copepods: morphology and function

Mate location and recognition are essentially asymmetrical processes in the reproductive biology of calanoid copepods with the active partner (the male) locating and catching the largely passive partner (the female). This behavioural asymmetry has led to the evolution of sexual dimorphism in copepods, playing many pivotal roles during the various successive phases of copulatory and post-copulatory behaviour. Sexually dimorphic appendages and structures are engaged in (1) mate recognition by the male; (2) capture of the female by the male; (3) transfer and attachment of a spermatophore to the female by the male; (4) removal of discharged spermatophore(s) by the female; and (5) fertilization and release of the eggs by the female. In many male calanoids, the antennulary chemosensory system is enhanced at the final moult and this enhancement appears to be strongly linked to their mate-locating role, i.e. detection of sex pheromones released by the female. It can be extreme in calanoids inhabiting oceanic waters, taking the form of a doubling in the number of aesthetascs on almost every segment, and is less expressed in forms residing in turbulent, neritic waters. Mate recognition is a process where chemoreception and mechanoreception presumably work in conjunction. The less elaborate male chemosensory system in the Centropagoidea is counterbalanced by females playing a more active role in generating hydromechanical cues. This is reflected in females in the shape of the posterior prosomal margin, the complexity of urosomal morphology and the size of the caudal setae. Visual mate recognition may be important in the Pontellidae, which typically show sexual dimorphism in eye design. The most distinctive sexual dimorphism is the atrophy of the mouthparts of non-feeding males, illustrating how copepod detection systems can be shifted to a new modality at the final moult. In the next phase, the male captures the female using the geniculate antennule and/or other appendages. Three types of antennulary geniculations are recognized, and their detailed morphology suggests that they have originated independently. Grasping efficiency can be enhanced by the development of supplemental hinges. The scanty data on capture mechanisms in males lacking geniculate antennules are reviewed. It is suggested that the loss of the antennulary geniculation in many non-centropagoidean calanoids has evolved in response to increasing predator pressure imposed on pairs in amplexus. Spermatophore transfer and placement are generally accomplished by the modified leg 5 of the male. In some males, leg 5 consists of both a chelate grasping leg and a spermatophore-transferring leg, whereas in others, only the latter is developed. Tufts of fine setules/spinules and/or sclerotized elements on the terminal portion of the leg are involved in the transfer and attachment of the spermatophore. The configuration of gonopores, copulatory pores and their connecting ducts in the female genital double-somite is diversified in the early calanoid offshoots such as Arietellidae and Metridinidae, whereas in more derived groups, it is constant and invariable, with paired gonopores and copulatory pores located beneath a single genital operculum. The absence of seminal receptacles in most Centropagoidea limits the female's ability to store sufficient sperm for multiple egg batches, suggesting that repeated mating is necessary for sustained egg production. Discharged spermatophores are usually removed by the female leg 5 and/or specialized elements on other legs. In Tortanus (Atortus) Ohtsuka, which has rudimentary fifth legs in the female and complex coupling devices in the male, a spermatophore supposedly remains on the female urosome, since eggs appear to be released from a ventral opening of the spermatophore. The type of sexual dimorphism is closely related to habitat and biology. Some hyperbenthic families never show multiplication of aesthetascs on the male antennule, whereas families of the open pelagic realm such as the Aetideidae always have non-feeding males exhibiting secondary multiplication of antennulary aesthetascs. The various aspects and diversity of calanoid sexual dimorphism are herein considered in an evolutionary context.     

Seasonal variability in the individual carbon content of the calanoid copepod Acartia bifilosa from the northern Baltic Sea

The seasonal variation in the carbon content of the calanoid copepod Acartia bifilosa was studied in the northern Baltic Sea. Monthly length-weight relationships were followed from November 1990 to October 1991 by analysing the organic carbon content of individuals, using a high temperature combustion method. The monthly length (L) on carbon (C) regressions of copepodites were best described by power functions (C = aL^b), where 75% to 88% of the variation could be explained by length. Nauplii length explained less of their carbon content (63–71 %). The carbon content per length was highest in summer when the nutritional situation was good and temperature at its highest. However, the carbon-length regressions differed significantly between months, except in winter, when two subsequent months had similar regressions. The results pointed out the importance of seasonal variation as well as the risk of making errors if biomass estimates based on length on carbon regressions are used. Carbon analysis should, if possible, be done on every sample or should at least be tested as to whether the regressions are consistent with the study material before estimations are calculated.     

Long-term dynamics of main mesozooplankton species in the central Baltic Sea

Long-term dynamics (1959–1997) of the copepod speciesPseudocalanus elongatus, Temora longicornis, Acartia spp. andCentropages hamatus, as well as the taxonomic group of cladocerans,are described for the open sea areas of the central Baltic Sea.Differences between areas, i.e. Bornholm Basin, Gdansk Deepand Gotland Basin, as well as between 5 year periods, were investigatedby means of Analysis of Variance (ANOVA). No significant differencesin mesozooplankton biomass between areas were found. On theother hand, clear time-trends could be demonstrated and relatedto salinity and temperature, with P.elongatus biomass mainlydependent on salinity and T.longicornis, Acartia spp. and cladoceransbiomasses dependent, to a large extent, on thermal conditions.Decreasing salinities since the early 1980s due to a lack ofmajor inflows of highly saline water from the North Sea andincreased river run-off, both triggered by meteorological conditions,obviously caused a decrease in biomass of P.elongatus. Contrarily,the standing stocks of the other abundant copepod species andcladocerans followed, to a large degree, the temperature developmentand showed, in general, an increase. The shift in species compositionduring this period is considered to be a reason for decreasinggrowth rates of Baltic herring (Clupea harengus) since the early1980s, and for sprat (Sprattus sprattus) since the early 1990s.Generally, it is suggested that low mesozooplankton biomassesin the 1990s were caused, at least partially, by amplified predationby clupeid fish stocks.     

Mechanoreceptors in calanoid copepods: designed for high sensitivity

The mechanoreceptors of the first antennae of Pleuromamma xiphias, a mesopelagic calanoid copepod, are critical for the detection of potential threats. These receptors exceed the physiological performance of other crustacean mechanoreceptors in sensitivity to water velocities as well as in frequency response. A study of these receptors was initiated to elucidate structure–function relationships. Morphologically, the receptors resemble the arthropod scolopidial organs by the presence of a scolopale tube. However, the rigidity of the copepod receptors greatly exceeds those described for crustaceans and other arthropods. The scolopale tube completely encloses the distal dendrites and it is firmly anchored to the cuticle. Microtubules are organized in register and arise from microtubule subfibers associated with crescent-shaped rods which extend from the basal body region to the setal socket. The distal dendrites are filled with a large number of cross-linked microtubules. Termination of the distal dendrites inside the lumen of the setae is gradual with a firm anchoring to the cuticle. A likely mechanism for mechanotransduction would involve a linkage between individual microtubules and mechano-gated channels in the dendritic membrane. The rigidity probably contributes to the high frequency sensitivity, and termination of the dendrite inside the seta increases the overall sensitivity of these receptors.     

Spatio-temporal variation in stable isotope and elemental composition of key-species reflect environmental changes in the Baltic Sea

Biogeochemistry - Carbon and nitrogen stable isotope ratios are increasingly used to study long-term change in food web structure and nutrient cycling. We retrospectively analyse elemental...