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).     

Trophodynamics and seasonal cycle of the copepod Pseudocalanus acuspes in the Central Baltic Sea (Bornholm Basin): evidence from lipid composition

Seasonal lipid dynamics of the copepod Pseudocalanus acuspes were studied in the Bornholm Basin (Central Baltic Sea) on a monthly basis from March 2002 until March 2003 and were interpreted in light of life cycle strategies and diet selection. The individual total lipid content of females ranged from 0.9 to 1.8 μg, with relative wax ester content reaching a significant maximum in May (44% of total lipids) and minimum (17% of total lipids) in April and November. Significant changes in size, lipid content, lipid classes and fatty acid composition of structural as well as storage lipids suggested five characteristic seasonal phases that were induced by different feeding histories and environmental conditions. Storage lipids were characterized by 18:1(n−9) as major component, which ranged between 44% of total fatty acids in June and 23% in February. The strong coherence between 18:1(n−9) in the seston lipids and the occurrence of ciliates emphasized the importance of ciliates in the diet of P. acuspes. As indicated by changes in the amounts of fatty acid markers, other food sources varied over the year, suggesting an opportunistic feeding behavior. The spring period was characterized by an increase in typical diatom and dinoflagellate markers, whereas other sources, potentially cyanobacteria, became more important during summer. The life cycle strategy is discussed with respect to extant adaptations to high latitudinal habitats.     

Seasonal variations in distribution and population structure of Microcalanus pygmaeus and Ctenocalanus citer (Copepoda: Calanoida) in the eastern Weddell Sea,Antarctica

The abundance, vertical distribution and population structure of two important small calanoid copepod species, Microcalanus pygmaeus (G. O. Sars) and Ctenocalanus citer Heron and Bowman, were studied in the eastern Weddell Sea in summer (January/February 1985), in late winter/early spring (October/November 1986) and in autumn (April/May 1992). The population of Microcalanus pygmaeus consisted mainly of copepodite stages CII and CIII in late winter/early spring and were concentrated between 500 and 200 m depth. In summer, stage CIV was the modal stage and the bulk of the population had ascended above 300 m. In autumn the population structure was bimodal with CI and CV dominating. Most of the population was concentrated between 300 and 200 m. In all investigation periods M. pygmaeus had their maximal concentrations in the thermo-pycnocline. The developmental stages CIII to CV of Ctenocalanus citer formed the bulk of the population in late winter/early spring. In October all developmental stages had their main distribution between 500 and 200 m, except females, which were concentrated in the upper 50 m. In November most of the population occurred between 200 and 50 m. The summer population was concentrated in the upper 50 m, and numbers increased dramatically as the new cohort hatched. Copepodite stages CII and CIII dominated the population at the end of January, while CIV dominated 2 wk later. In autumn, CV was the modal stage. The majority of the population was concentrated in the upper 100 m, but there was an increase in abundance below 300 m compared to summer. Age structure changed with depth with a younger surface population and an older one in deeper water layers. The seasonal change in number of M. pygmaeus is much smaller than that of C. citer; the summer:winter:autumn ratio of the former being about one, whereas the winter:summer/autumn of the latter was about nine. Early copepodite stages and adults of M. pygmaeus occurred throughout all investigation periods. The large proportion of early copepodite stages in April and in mid-October suggests autumn and early to midwinter breeding. Apparently, M. pygmaeus may reproduce and grow year-round or perhaps has a 2-yr life-cycle. In contrast, the dramatic increase in abundance of early copepodite stages of C. citer in summer suggests springtime reproduction.     

Life cycles ofCalanoides acutus,Calanus simillimus andRhincalanus gigas (Copepoda: Calanoida) within the Scotia Sea

The life cycles and distribution of three dominant copepods,Calanoides acutus, Calanus simillimus andRhincalanus gigas were studied from the Discovery collections in the Scotia Sea earlier this century.C. simillimus is a sub-Antarctic species which mates in the top 250 m mainly in spring. The rapid development of the summer generation may allow a second mating period and a smaller second generation to appear in late summer.C. simillimus remains in the surface layers for a longer period thanCalanoides acutus orR. gigas, and its depth distribution is bimodal throughout the winter.R. gigas is most abundant in sub-Antarctic waters to the north of the Polar Front. It mates within the top 750 m later in spring, and development seems less synchronised than that of the other two species, with egg laying and the growth season being more protracted. Stages CIII and CIV are reached by the first autumn and further development resumes very early the following spring. It is not clear whether the majority then spawn or whether a further year may be needed to complete the life cycle. The predominantly Antarctic species,C. acutus mates below 750 m in middle to late winter and the summer generation develops rapidly to either CIV or CV. Its lifespan seems typically 1 yr, but some of the CVs which fail to moult and spawn in winter survive into their second summer, and their subsequent fate is uncertain. The cold-water speciesCalanus propinquus is comparatively rare in the Scotia Sea and aspects of its distribution and life cycle are briefly described for comparison. Regional variations in the timing of these events were apparent forC. simillimus and possiblyCalanoides acutus, but were not seen inR. gigas. Their geographic and vertical separation, together with their asynchronous life cycles support the concept of habitat-partitioning of these dominant herbivores.     

Distribution and relative abundance of Pseudocalanus moultoni and P. newmani (Copepoda: Calanoida) on Georges Bank using molecular identification of sibling species

The sibling species, Pseudocalanus moultoni (Frost, 1989) and P. newmani (Frost, 1989), occur sympatrically on Georges Bank. Taxonomic discrimination of the species relies on subtle morphological characteristics, making routine identification of the species very difficult. DNA sequence variation of two mitochondrial genes, 16S rRNA and cytochrome oxidase I (COI), reliably discriminated P. moultoni and P.␣newmani. Levels of DNA sequence variation for both genes were consistent with those between species of calanoid copepods. A molecular systematic protocol (based on allele-specific PCR amplification) was designed from the COI sequences and used to discriminate females of the two species. The distributions and relative abundances of the two species were mapped for April 1996 based on samples of 15 to 30 females from 12 collections across Georges Bank. The results of this study indicated that P. moultoni females predominated along the northern flank of Georges Bank, while P. newmani females were common on the southern flank, deeper than the 60-m isobath. Received: 16 May 1997 / Accepted: 26 April 1998     

Female genital structures of Metridinidae (Copepoda: Calanoida)

The female genital structures of six calanoid copepod species, belonging to the genera Gaussia, Metridia and Pleuromamma, were studied using light and scanning electron microscopy (SEM). The copulatory pores and seminal receptacles are paired in Gaussia and Metridia, but unpaired in Pleuromamma. A thin epicuticle and a spermatophoral plug are the mechanisms by which the pores are closed before and after copulation. The pores open directly into the receptacles, which are reduced to shallow integumentary depressions in Gaussia. The mode of insemination suggests two matings in females of Gaussia and Metridia, but only one in Pleuromamma. Paired gonopores and egg-laying ducts are present in the three genera, with a characteristic, closed semicircular configuration; these are opened during egg-laying by the action of retractor muscles of the gonoporal plates. The seminal ducts, which open into the gonopores, are long, thin and paired in Gaussia and Metridia, whereas in Pleuromamma they are short, broad and unpaired. The opening of the seminal and egg-laying ducts is synchronized. The shell ducts are paired in Gaussia and Metridia, unpaired in Pleuromamma; these arise from glands situated in the lateral expansions of the last prosomite and lead into the distal part of each egg-laying duct in Gaussia and Metridia and of the egg-laying duct receiving the seminal duct in Pleuromamma. The position and anatomy of these structures are compared to those of other families and genera, and a functional interpretation of their morphology is proposed. The main evolutionary trends of the different structural patterns of female calanoid genitalia are presented in tabular form. Received: 30 December 1996 / Accepted: 11 February 1997     

Mating behavior of Centropages typicus (Copepoda: Calanoida)

Observations were made on approximately 300 mating couples of Centropages typicus Krøyer. Mating behavior consists of 4 distinct stages: pre-copulation, spermatophore extrusion, copulation, and post-copulation. Successful copulation requires proper positioning of the spermatophore complex on the female urosome and is dependent upon precise movements and orientation of the male during the mating encounter. Sub-optimal mating encounters, resulting in the improper placement of the spermatophore complex, were observed and adaptations tending to reduce spermatophore wastage are discussed.Contribution No. 62, Harbor Branch Foundation, Inc.     

Mating behavior of Labidocera aestiva (Copepoda: Calanoida)

The calanoid copepod Labidocera aestiva Wheeler exhibits a ritualized mating behavior. Sensory hairs and spines and integumental pores on the male and female are believed to play a regulatory role in the mating encounter. Scanning electron microscopy (SEM) revealed approximately 20 unusual pore structures (pit-pores) on the right ventrolateral surface of the female's abdominal segment, the area where the spermatophore coupling device is attached. Prior to spermatophore transfer, the male vigorously strokes this area with the modified endopod of his left fifth leg to prepare the pit-pore region for attachment of a new spermatophore. Transmission electron microscopy (TEM) indicated that the cells underlying each pit-pore may have a secretory function. We hypothesize that the pit-pores secrete a substance which dissolves the cement binding the coupling device to the abdomen and aids the female to remove a discharged spermatophore from her urosome.Contribution No. 116, Harbor Branch Foundation, Inc.     

Feeding of Clausocalanids (Calanoida,Copepoda) on naturally occurring particles in the northern Gulf of Aqaba (Red Sea)

A total of 12 feeding experiments were conducted in the northern Gulf of Aqaba during spring (March/April) and autumn (September/October) 2002 at the Marine Science Station (MSS) in Aqaba. Females of three species of clausocalanids were selected: Clausocalanus farrani, C. furcatus and Ctenocalanus vanus. Natural occurring particle (NOP) larger than 5 μm were investigated as food source. The ambient chlorophyll a concentration at sampling depth (∼70 m) ranged between 0.15 and 1.00 μg chl a l⁻¹ and NOP concentrations ranged between 1.78 and 14.0 × 10³ cells l⁻¹ during the sampling periods. The division of particles into five size classes (5–10, 10–20, 20–50, 50–100 and >100 μm) revealed that most of the particles were found in the size classes below 50 μm (81–98%), while most of the natural occurring carbon (NOC) was concentrated in the size classes larger than 20 μm (70–95%). Ingestion rates were food density dependent rather than size dependent ranging between 0.02 and 1.65 × 10³ NOP ind⁻¹ day⁻¹ and 0.01 and 0.41 μg NOC ind⁻¹ day⁻¹, respectively, equivalent to a body carbon (BC) uptake between 0.4 and 51.8% BC day⁻¹. The share of the size classes to the total ingestion resembled in most cases the size class composition of the natural particle community.     

Mating behavior in three species of Pseudodiaptomus (Copepoda: Calanoida)

Quantitative studies of mating behavior between conspecific and heterospecific pairs of three closely related species in the genus Pseudodiaptomus reveal differences in the timing of life-history events, species-specific attraction of mates, distinctive tactile cues, multiple spermatophore attachments, and gametic incomptibility or hybrid inviability. These differences may function as reproductive isolating mechanisms among calanoid copepods. Differences in breeding behaviors and associated morphological features are greater between sympatric than between allopatric species, indicating that selection reinforces both behavioral and morphological divergences.     

Nutrition studies in the nauplius larva of Calanus pacificus (Copepoda: Calanoida)

Nauplii of Calanus pacificus were raised on a mixture of algae. Details of the mouth-parts, such as denticles, labial palps and lobes, setations and structure of the masticatory teeth were examined by scanning electron microscopy (SEM). Under the experimental conditions (15°C and 300 gC l^-1), exponential growth coefficients for the period Nauplii II–VI were 0.179 for carbon and 0.228 for nitrogen. C:N ratios dropping from 5.1 to 4.7. Growth was isochronous, each stage lasting 1.5 days. Respiratory losses were 15 to 19.6% of body carbon daily. Nauplii raised on a given alga showed higher rates of ingestion in the presence of this food, compared to nauplii switched to other algae. Minimal threshold concentrations for feeding were found, depending on the size of the food offered and ranging from 5.8 gC I^-1 for Lauderia borealis (28.7 m spherical diameter) to 47.1 gC 1^-1 for Chlamydomonas sp. (11.0 m). Unlike the Copepodite I stage, Nauplii II–VI larvae were not able to ingest small cells such as Isochrysis galbana (4.3 m), or very large ones such as Ditylum brightwellii (47.5 m) at more than maintenance rations. Below the critical concentration for maximal feeding, ingestion was clearly dependent on size of the cells offered, but the size-dependent relationship was different for diatoms and non-diatoms. Filtering rates increased from a threshold concentration to a maximal rate at about 50 gC 1^-1, and decreased at higher concentrations. Critical concentrations ranged from 125 gC 1^-1 for L. borealis to 1000 gC 1^-1 for Chlamydomonas sp. Maximal daily rations ranged between 100 and 150% of body carbon.     

Calanoid resting eggs in the Baltic Sea: implications for the population dynamics of Acartia bifilosa (Copepoda)

Resting eggs of Acartia spp. (presumably A. bifilosa Giesbr.) at densities between 65 and 125 eggs cm^-2 were found in sediment off the southern coast of Finland and represent the first observations of dormant calanoid eggs in the Baltic Sea. The highest egg densities were found at sediment depths between 4 and 8 cm. The hatching percentage varied between 0 and 90% in the different sediment layers and at different experimental temperatures (13, 16 and 19°C). Some nauplii emerged from the eggs extracted from sediment layers 9 to 10 cm beneath the surface layer and other nauplii emerged from eggs after 82 d of dark storage at 3°C. Light was not needed to trigger hatching; the nauplii were able to hatch in darkness when the eggs were resuspended in filtered seawater. Indirect evidence suggests that the marked spring peak of Acartia spp. nauplii seen in a monitoring data from the years 1973 to 1984 was derived from benthic eggs and not from spawning. The possible mechanisms for induction of hatching and the adaptive benefits of resting egg production for A. bifilosa are discussed.     

Tisbe (Copepoda: Harpacticoida) species from the Lagoon of Venice. I. Seasonal fluctuations and ecology

Distribution and abundance of Tisbe species were studied throughout a period of 3 years at 3 stations in the Lagoon of Venice (Italy). At 2 other stations samples were taken occasionally, as also at the Lido station (open sea). Twelve species may be considered as common inhabitants of the lagoon, and 3 species appear to be occasional transients, compared to the 9 species found at the Lido. A comparison of the physical data shows that the stations were similar in temperature and salinity, but differed primarily in pH values, sediment characteristics, algal substratum and faunal benthic community. The fact that remarkable differences in species composition and distribution of Tisbe could be observed between the various parts of the lagoon and the lagoon and the open sea, indicates that temperature and salinity are not so important for species diversity as are biotic factors. A particularly strong difference existed between samples taken at the bottom and from pilings, which constitute a very special biotope for the benthic communities in the lagoon and seem to be a preferred habitat for T. lagunaris and T. cucumariae. The data suggest that there is a certain interaction between T. holothuriae and the other species. Whenever T. holothuriae was abundant, the other species were rare, but T. clodiensis, T. dobzhanskii (Stations 1 and 2) and Tisbe sp. (Station 3) attained high relative abundance whenever T. holothuriae decreased considerably in number. T. holothuriae represents certainly the hardiest species of the lagoon, displaying a great ability to cope with stress conditions, even those present in the interior lagoon where strong fluctuations in salinity, temperature, pH and oxygen prevail.     

Suspected mechano- and chemosensory structures of Temora longicornis (Copepoda: Calanoida)

The antennulary (A1) setae of Temora longicornis Müller are suspected of being mechanosensory. In the present study, the fine structure of the antennule, setae and pegs was examined by scanning and transmission electron microscopy. Two setae arise proximal to each antennule segment junction. The supraaxial (Type 1) seta of each pair is innervated by a pair of ciliary dendrites and has the morphological characteristics of a mechanoreceptor. The sub-axial (Type 2) seta may be chemosensory, and is innervated by a varying number of dendrites, but always more than two. The antennulary pegs do not appear to be innervated. T. longicornis is directionally sensitive to water disturbances, and this is consistent with the physical structure of the setae. Integumental structures on the body and other non-feeding appendages are described. These observations support previous behavioural experiments which suggested the antennule to be the major site for mechanoreception in copepods; however, mechanoreceptors must also occur on other parts of the body.     

Community interactions and zoogeography of the Indian Ocean Candaciidae (Copepoda: Calanoida)

Eighteen species of Candaciidae have been identified from collections made at 339 stations in the Indian Ocean. Most species are zonally distributed; however, on the eastern and western sides of the Indian Ocean, species ranges are extended north or south by boundary currents. Factor analysis was used to cluster phenotypically similar species based on 130 characters taken from the maxilla (a feeding appendage), the first swimming foot, and the fifth foot (a secondary sexual structure). Four morphological clusters were extracted. Clusterings based on separate factor analyses limited to characters from feeding or from sexually adapted appendages are in substantial agreement with the clusters based on composite morphology. Two geographically recurrent groups of species were also identified, one from equatorial waters, one from the central gyre. Each recurrent group is composed of species exceptionally different in body size and belonging to different morphological clusters. Niches of the species are compared using Hutchinson's multidimensional hypervolume as a model. An attempt is made to describe the niche of each species in terms of environmental variables measured at stations where the species was abundant. Environmental variables (dimensions) included in this study are space, time, temperature, food, oxygen and salinity. Mean niche adaptations of most of the species are separable from congeners along at least one niche dimension. It is proposed that community relationships among the Candaciidae developed to prevent both gamete loss and to avoid trophic competition. It is postulated that newly evolving species underwent contemporaneous displacement of both secondary sexual and trophic characters as a condition for sympatry, or diverged in their physiological adaptations to escape sympatric interference.Contribution No. 3686 of the Woods Hole Oceanographic Institution. This research was supported by NSF Grants GB 27405 and GA 43126 and is based on a doctoral dissertation submitted to the Curriculum in Marine Sciences of the University of North Carolina at Chapel Hill.     

Relationships between RNA content and egg production rate in Acartia bifilosa (Copepoda, Calanoida) of different spatial and temporal origin

To evaluate factors regulating RNA content (RNA, μg RNA female⁻¹) – egg production rate (EPR, eggs female⁻¹ day⁻¹) relationship and to develop a model for in situ egg production rate estimates for Acartia bifilosa, we (1) measured EPR and RNA in females sampled at geographically distant areas at varying temperature (T, °C), (2) determined environmental (station, season, and T), endogenous (prosome length (PL), mm and EPR) variables that influence RNA levels, and (3) explored a set of multiple regression models to predict EPR from RNA, PL, station, season, and T. Egg production experiments were carried out in spring and summer 2005 in the Gulf of Finland and in the western part of the northern Baltic proper. We found that up to 88% of the RNA variation could be explained by variations in PL, EPR, and season/T. In explaining the RNA variability, PL played a major role followed in order of importance by EPR and season/T. Further, PL, RNA, and season/T explained up to 53% of the variation in EPR, nearly half of which is explained by RNA alone. The effect of spatial origin was never significant, suggesting that the derived relationships are general for A. bifilosa inhabiting northern Baltic proper. An erratum to this article is available at .     

Vertical and spatial distribution of biogenic silica in the sediment of the gulf of Riga,Baltic Sea

The vertical and spatial concentration trends of biogenic silica (BSi) were investigated in a variety of sediment types from the Gulf of Riga. These trends were compared with the concentration of organic carbon to examine how variations in the mass sedimentation and the dissolution rate of BSi versus bacterially mediated oxidation of organic matter affect the distribution of BSi in different environments. The effects of important physical and biogeochemical factors on the distribution of BSi have also been examined.

The concentration of BSi in the surface sediment at accumulation bottoms ranges between 0.8–1.5 mmol/g with an average concentration of 1.1 mmol/g. This is about two times the concentration that has been reported in similar sediments in the open Baltic proper. Much lower concentrations were measured in erosion and transitional areas; on average around 0.2 and 0.6 mmol/g, respectively.

A comparison between recent sedimentation rate measurements (²¹⁰Pb) and concentrations of BSi within different accumulation bottoms suggests an approximate inverse relationship with each other. The importance of variation in the sedimentation rate on the surface BSi concentration is supported by the fact that higher C_org/BSi ratios are found in periphery accumulation zones close to transition areas (larger supply of carbon rich allochthonous material that dilutes settled amounts of BSi) than at accumulation bottoms in the central part of the Gulf. The C_org/BSi ratio in the sediment is always lower than in living siliceous algae which shows that the microbial degradation of organic carbon is always greater than the inorganic dissolution of BSi, irrespective of the assemblage of silica and non‐silica species.

The general vertical concentration pattern at accumulation sites showed a decreased concentration with increased depth down to 2–6 cm below seafloor where the concentration gradually equalizes. The concentration in the equalizing zone is on average about 20% lower than in the uppermost centimeter. This implies that the main part of the BSi entering the accumulation bottoms is permanently buried and that a great amount of silica already has been released during halmyrolysis. The spatial and vertical BSi trends at sites in transitional and erosional areas reflects prevailing sedimentation conditions with a low concentration in sections with a dominance of sand and gravel (no deposition of fine material) and a higher concentration in the sections containing stiff glacial clay (former accumulation area).

The average annual burial amount of BSi in the Gulf has been estimated to be around 50 000 tons of which most is buried in accumulation bottoms outside the central muddy part of the Gulf. The buried amount is about 60% of the annual river load of silica.     

Fecundity studies on Acartia tonsa (Copepoda: Calanoida) in standardized culture

Acartia tonsa Dana longevity and egg production data were studied over a 4-year period under standardized culture conditions. Egg-laying patterns and daily rates were evaluated as bioassay tools for measuring the nutritional value of various copepod foods, as well as assessing other aspects of environmental quality. Altogether, 337 females were observed in various aspects of this work. Experiments were performed with newly matured females in loosely covered crystallizing dishes. Fresh water and a standard concentration of algae were supplied daily, and a running daily tally was kept of the eggs produced. The effects of male copepods on egg production were analyzed. Males were either removed after the first eggs were observed or they remained with the female throughout her life span. Daily egg-laying rates of this latter group of females appeared to be more stable, but abnormal embryo development occurred if the males were not removed after fertilization. Likewise, the results of females which were not mated were studied. Life spans were shorter and egg production rates were lower than those of contemporary mated females. Unmated females were subjectively observed to be less active than their mated counterparts. The effects of continuous and temporary starvation on egg-laying were also observed. Under both normal and starved conditions, males appeared to have shorter life spans than either mated or unmated females. During starvation there were no real differences between the life spans or egg productions of mated or unmated females. Non-nutritional particulate matter was evaluated as a food source. It was ingested by the females, but did nothing to prolong the egg-laying period. Finally, the three algal components of the standard ration were evaluated. Females produced equally well with a mixed diet or with Thalassiosira pseudonana (100 m³) or Chroomonas salina (140 m³), the two larger species of this diet, but when Isochrysis galbana (40 m³) was fed, females reacted as if in a starved situation.