Diel vertical migration and feeding of copepods at an oceanic site near South Georgia

Twelve Longhurst Hardy Plankton Recorder (LHPR) profiles were taken over a 16 h period in January 1990, in order to study feeding of four copepod species at an Antarctic oceanic site near South Georgia. Vertical distributions of their life stages, as well as those of dominant competitors and predators, are described in relation to the feeding cycles of Calanoides acutus CV, Calanus simillimus CV, Calanus propinquus CV and Rhincalanus gigas CIII, CV and CVI. Comparisons with vertical ring-net catches, which were used for concomitant gutevacuation experiments, demonstrated the suitability of the LHPR for these fine-scale studies. Planktonic predators, with the exception of the diel migrant Themisto gaudichaudii, resided deeper than the herbivores. During the day and around midnight, when feeding rates were low, species and stages reached their maximum vertical separation. At these times, new generation copepodites of the four species lived progressively deeper and the overwintered generation (i.e., R. gigas Stages CIV, CV, CVI) were progressively shallower. During the afternoon or evening (depending on species), all stages older than CII, as well as Euphausia frigida and T. gaudichaudii, migrated upwards, to amass in the surface mixed layer. Feeding was restricted to darkness, although R. gigas commenced several hours before dusk. In detail their migration and feeding differed widely, with combinations of unimodal and apparent bimodal cycles. As a whole, the results suggest that (1) feeding could occur during sinking as well as during upward migrations, (2) upward migrations were not always associated with feeding increases, and (3) individuals appeared to descend after filling their guts.     

Phosphorus dynamics associated with phytoplankton blooms in eutrophic Mikawa Bay,Japan

Changes in the phosphorus components of the particulate matter in seawater were studied in the eutrophicated waters of Mikawa Bay, Japan, during summer 1981. The contents of particulate phosphorus and hot-water extractable intracellular phosphorus displayed remarkable changes associated with phytoplankton blooms caused by wind-induced or upwelling-associated nutrient enrichment from the bottom water layers. Nanoplankton <10 m accounted for much of the particulate phosphorus (70 to 79% in June and July, and 44 to 78% in August and September); the contribution of large-sized phytoplankton >25 m varied from 9 to 49%, the peak values being attained under red-tide conditions. The capacity for phosphorus storage in cells was low in nanoplankton cells, high in large phytoplankton species. Differences in rates of phosphorus storage and growth between nanoplankton and large phytoplankton accounted for fluctuations in particulate phosphorus which were closely associated with fluctuations in phytoplankton blooms in Mikaw Bay.     

Size-weight relationships and biomass of tropical neritic copepods off Kingston,Jamaica

Nine genera of neritic tropical copepods were collected near Lime Cay, Jamaica, between July 1985 and January 1987. Length-weight regressions were derived for each genus (R ²=0.74 to 0.98), for all calanoids combined (R ²=0.88), and all cyclopoids combined (R ²=0.85). Width-weight regressions were also derived for the same genera but coefficients of determination were generally lower (R ²=0.52 to 0.98) and were much lower forOithona spp. (R ²=0.21). Over a 12 mo period, biomass estimates generated from these length-weight regressions differed by only 3% from direct weight determinations. There were no significant annual variations in prosome lengths for copepodite Stages 1 to 5 inCentropages velificatus, Paracalanus aculeatus orTemora turbinata; prosome lengths in femaleT. turbinata did significantly vary seasonally. No significant seasonal differences in length-weight relationships were observed forC. velificatus. The mean ash content of mixed copepod samples was 6.4%, and the energy density was 25.0 kJ g^–1 AFDW. No significant loss of weight was observed 10 mo after preservation in 10% formalin.     

Spatio-temporal changes in diversity and community structure of planktonic copepods in Sagami Bay, Japan

Seasonal changes in diversity and community structure of planktonic copepods at a shelf site in Sagami Bay, Japan was studied in relation to cross-shelf interaction of species components. Seasonal mesozooplankton samples were collected from the shelf station (St. M) of the north-west part of Sagami Bay from 1995 to 1997. Vertical multi-layered samples were collected near the center of Sagami Bay (St. P) in June 1996. A total 185 copepod species were identified from the two stations. We observed a clear seasonal succession in calanoid diversity and community structure at St. M from a simple shelf water community (>11 species) during spring blooming periods to highly diverse and mixed communities (ca 20–30 species) of shelf water species coupled with various Kuroshio Current species during late summer to autumn. Cluster and non-metric multidimensional scaling ordination analyses showed two distinct calanoid community groups. One group, which included samples of St. M and the surface layer of St. P, consisted of shelf water species, such as Calanus sinicus, Ctenocalanus vanus, Paracalanus spp., and Kuroshio species, such as, Canthocalanus pauper, Scolecithrix danae, etc. The other cluster was restricted to the samples collected from mid and deep layers at St. P, which consisted of meso- and bathypelagic species and Oyashio species (cold-current species, such as Neocalanus cristatus, Pseudocalanus spp., Eucalanus bungii and Metridia pacifica). In the mid and deep layers at St. P, the population of dormant copepodid stage V (CV) of Eucalanus californicus and C. sinicus were dominant. The deep CV population of C. sinicus might be ecologically discriminated from the surface and shelf water population due to their larger body length and dormant life cycle. E. californicus was also collected at the shelf site during each spring bloom period, whereas the population might descend into the mid- and deep-layers of the central bay before summer. Our results suggest that the seasonal fluctuation of community structure in the shelf water was controlled by both physical (Kuroshio Current) and biological factors, i.e., spring bloom and ontogenetic vertical migration of E. californicus. In particular, transport and diffusion processes of Kuroshio Current in Sagami Bay played a key role in controlling the shelf water calanoid community.     

Water-soluble vitamins in natural plankton (copepods) during two consecutive spring blooms compared to vitamins in Artemia franciscana nauplii and metanauplii

Weight-specific contents of asorbic acid, thiamin, riboflavin, niacin, pantothenic acid, vitamin B₆, biotin, folate and vitamin B₁₂ were analysed in natural zooplankton collected from a sea water lagoon in a halibut fry production plant. The samples of natural zooplankton were collected from April/May to the end of June during two consecutive spring blooms. For comparison, the same vitamins were measured in nauplii of Artemia franciscana, just after hatching and after enrichment for 24 and 48 h with a commercial diet. The changes in vitamin content in zooplankton, which consisted mainly of Temora longicornis, during the two spring blooms were <2.5-fold for each specific vitamin. The large number of high quality halibut fry produced using the plankton as a dietary source suggests that the plankton contained sufficient amounts of all water-soluble vitamins. With the exception of thiamin and ascorbic acid, Artemia– both newly hatched and enriched for 24 and 48 h – contained higher levels of water-soluble vitamins than natural zooplankton. Enrichment feeding of Artemia for 48 h stabilised or increased the level of the water-soluble vitamins. Received: 8 March 1999 / Accepted: 21 February 2000     

Abundances,growth rates,and production of tropical neritic copepods off Kingston,Jamaica

Weekly samples were collected near Kingston, Jamaica in 27 m vertical hauls, using 200 and 64µm mesh plankton nets, from July 1985 to January 1987. Thirtytwo copepod species were identified; nauplii and all copepodite stages were enumerated. Total copepod abundance ranged from 2.56 to 87.3 × 10⁴ m^–2. The annual abundance cycle was bimodal with peaks in October–November and May–June corresponding to the rainy seasons. Mean annual copepodite biomass was 0.15 g AFDW m^–2 ranging from 0.03 to 0.41 g AFDW m^–2. Mean generation time (from egg to adult) at 28°C was 19.5 d for the common speciesCentropages velificatus, Paracalanus aculeatus, andTemora turbinata. Isochronal development was demonstrated for copepodites ofP. aculeatus andT. turbinata, but not forC. velificatus. Mean daily specific growth rates (G) were 0.63, 0.63, and 0.48 d^–1 forC. velificatus, P. aculeatus, andT. turbinata, respectively. In general, daily specific growth rates decreased in the later copepodite stages. Thus, it is postulated that growth of later stages and egg production may be food limited. Annual copepodite production was estimated as 419 kJ m^–2 yr^–1, while annual exuvial production and naupliar production were 35 and 50 kJ m^–2 yr^–1, respectively. Egg production was estimated as 44% (184 kJ m^–2 yr^–1) of the total copepodite production. Thus, mean total annual copepod production was 688 kJ m^–2 yr^–1. This estimate is within the range of copepod production estimates in coastal temperate regions.     

Temporal patterns in diversity and species composition of deep-sea benthic copepods in bathyal Sagami Bay,central Japan

Little is known about temporal changes in the diversity and species composition of deep-sea metazoan meiofauna and their relationships with changes in the food supply. Those changes were studied for benthic copepod assemblages based on 2-year time-series data at a bathyal site in Sagami Bay (1430 m depth), central Japan, where annual fluctuation in the abundance of benthic foraminiferans was previously observed. Species diversity of benthic copepods at the site was as high as, or slightly higher than, that observed at other deep-sea sites, but did not fluctuate temporally through the study period. Multivariate analyses did not reveal any clear seasonal or directional change occurring over the longer term in their species composition, although there was some consistent pattern. These results indicate a lack of, or only weak, seasonality in the diversity and species structure of the deep-sea benthic copepod assemblages, even though the fresh organic food supply fluctuates seasonally. They also suggest that there are differences between copepods and foraminiferans in the response to changes in environmental factors, and that spatial differences in the composition of copepod communities are greater than temporal ones at this deep-sea site.Communicated by T. Ikeda, Hakodate     

Egg production and feeding by copepods prior to the spring bloom of phytoplankton in Fram Strait,Greenland Sea

Reproductively activeCalanus hyperboreus (Krøyer) andC. glacialis Jaschnov were captured in the upper 100 m of Fram Strait (77° to 79°N) in late winter 1987. There was no evidence of a phytoplankton bloom; chlorophylla concentrations were uniformly low (<0.1 mg m^–3), and nitrate concentrations were uniformly high (>11.3 mg-at m^–3). Gut-fullness measurements indicated that females were ingesting very little. The maturation state of gonads of bothC. hyperboreus andC. glacialis indicated that 75% of females were in a ripe condition consistent with observed egg laying. The lipid content of females laying eggs was reduced in both species compared to that of females not laying eggs. InC. hyperboreus the reduction was 39% and inC. glacialis it was 44%. All the evidence suggests that bothC. hyperboreus andC. glacialis were laying eggs in late winter by using lipids stored previously; they were not relying on ambient concentrations of phytoplankton. The daily rate of egg laying byC. glacialis using lipids in late winter exceeded the rate reported for summer when ambient food supplies have been shown to be necessary. It is suggested that individuals, spawned well in advance of the spring bloom of phytoplankton, may comprise a major portion of the annual recruitment to the entire population ofC. glacialis in this area, and that their life cycle can be completed within 1 yr. NeitherMetridia longa (Lubbock) norC. finmarchicus (Gunnerus) laid eggs during this study.     

Nitrogen utilization during spring blooms of marine phytoplankton in Bedford Basin,Nova Scotia,Canada

Spring diatom flowerings in Bedford Basin, Nova Scotia, almost exhausted the available nitrogen supplies. The supply of nitrate appeared to regulate the magnitude of the bloom. Other nutrient supplies may also have been limiting. Certain alterations in quantities of major metabolites could be interpreted in terms of nitrogen limitation. Nitrate, (nitrite), and ammonium were generally utilized simultaneously by Thalassiosira nordenskioldii, the dominant bloom organism. This species did not effectively utilize urea in either laboratory culture or in the natural environment. T. fluviatilis simultaneously utilized all nitrogen sources provided in laboratory experiments simulating natural nutrient conditions.     

Daytime surface swarms of the euphausiid Thysanoessa inermis off the west coast of Hokkaido,northern Japan

Biological characteristics of the daytime swarming of the euphausiid Thysanoessa inermis (Krøyer) were examined off the west coast of Hokkaido in the northeast Sea of Japan between 1982 and 1987. Swarms were composed exclusively of fully mature individuals; males possessed spermatophores at the ejaculatory ducts and females had attached spermatophores at the thelycum. Females predominated in most swarms, while males or nearly balanced swarms were infrequently encountered. Fully mature females occurred in the late swarming season, usually mid-April. Examination of stomachs indicated that feeding activity was low during the daytime surface swarming behavior. The body sizes of swarming T. inermis occurring around Yagishiri Island were significantly larger than those found simultaneously off the Shakotan Peninsula. This suggests that the euphausiid populations of the two areas are different.     

Chlorophyll blooms and the spring/neap tidal cycle: observations at two stations on the coast of Connemara,Ireland

Water temperature, chlorophyll a and phaeophytin were recorded every 2 to 4 d, in order to describe short-term variability in phytoplankton blooms. Data were collected from May to September 1984 and March to September 1985 at shore stations on the coast of Connemara, western Ireland. Comparisons with offshore data suggest that the shore data is representative of coastal water. Chlorophyll a was highest at neap tides in spring, and at spring tides in early summer. In late summer, transient chlorophyll blooms occurred at neap tides, possibly due to a localized accumulation of flagellates in stable conditions.     

Mosaic horizontal distributions of three species of copepods in the subarctic Pacific during spring

 Continuous abundance estimates (510 m resolution) of the copepods Neocalanus cristatus, N. flemingeri and Metridia pacifica were obtained with an electronic particle counter along cruise tracks in the subarctic western North Pacific in spring. For all three species, the number of patches decreased exponentially with increasing patch size. Most patches (63 to 83%) were dominated by one species, and patches of the same species more closely spaced than patches of different species. The patches of M. pacifica tended to coexist with those of N. cristatus. In contrast, patches of N. flemingeri rarely co-occurred with those of other copepods. These patterns were more clearly observed in fine-scale observations with sampling intervals of <31 m. Coherence analysis of copepod species pairs showed no characteristic scale at 2 to 50 km wave lengths. At shorter wave lengths (<2 km), frequent positive correlations were observed between N. cristatus and M. pacifica. Thus, the distribution of copepods appears to be a mosaic assemblage of patches of each copepod species. These results suggest that copepods have a mechanism to form species-specific aggregations, and the aggregation and segregation processes are maintained at a scale of <2 km. Received: 24 February 1999 / Accepted: 25 April 2000     

Vertical distribution,diurnal and seasonal migration of copepods in Saronic Bay,Greece

The problem of vertical distribution and the movements of copepods was studied from several plankton samples collected by vertical hauls in the Saronic Bay, Greece during two cruises (8 to 17 August and 21 November to 1 December, 1969). The species, whose vertical movements were analyzed, were divided into 3 groups: (1) Those which perform diurnal vertical migration (generally psychrophilic species found in summer samples in deeper layers and in reduced numbers). In autumn, the number of specimens is generally increased and many individuals reach the surface at night. From surface hauls it is known that these species abound in night surface hauls during the cold period. (2) Those species which execute a seasonal vertical migration. These, too are, in general, psychrophilic and found in summer in the deep water layers. In autumn, the population of the upper layers increases. From surface hauls it is known that these species abound in day and night surface samples. (3) Copepods which remained at the surface layer in both seasons of our cruises. These species are thermophilic and are absent from surface hauls during the cold period.     

In situ feeding rhythms of herbivorous copepods,and the effect of starvation

In situ diel variations in gut pigment contents of neritic (Acartia omorii andPseudocalanus minutus) and oceanic copepods (Calanus plumchrus andC. cristatus) were analyzed.A. omorii andP. minutus were sampled in Onagawa Bay on the east coast of Japan in May and August 1987, andC. plumchrus andC. cristatus were sampled in the Bering Sea in June 1986. Gut pigments were generally high at night, and bimodal feeding rhythms were observed in all species. The first peak of gut pigments occurred between sunset and midnight and was followed by a midnight decrease in gut pigment levels, resulting in eventual evacuation of the gut. The second peak was observed a few hours after sunrise. Incubation experiments indicated that ingestion rates of starved copepods were higher than those of acclimated copepods. This phenomenon was most notable at high food concentrations. Gut pigments of starved copepods rapidly increased after exposure to high concentrations of culturedThalassiosira decipiens. These findings suggest that in situ feeding behavior of herbivorous copepods includes periods of cessation or reduction in feeding during the night, and consequently, feeding activity is periodically enhanced with starvation. Starvation enhanced feeding behavior is most obvious in the large oceanic species,C. plumchrus andC. cristatus and is not distinct in small coastal species such asA. omorii.     

Annual population development and production by small copepods in Disko Bay,western Greenland

The population of small copepod species (approximately <1 mm) were investigated during an annual cycle in Disko Bay, western Greenland. The small species considered were Acartia longiremis, Pseudocalanus spp., Oithona spp., Oncaea spp., Microsetella spp., and Microcalanus spp. Most of the small species were present in the surface waters year round and numerically dominated the community, and in biomass from late summer and throughout winter. Oithona spp. was numerically the main contributor, while Pseudocalanus spp. dominated in terms of biomass. In the uppermost 50 m, maximum abundance, biomass and secondary production were observed in late September after the phytoplankton production practically had terminated and the winter initiated. The free spawning Acartia longiremis showed a strong seasonal fluctuation in biomass and egg production, in contrast to the egg carrying species Pseudocalanus spp. and Oithona spp. These had a long spawning season and maintained a more stable biomass year round. Secondary production was estimated by three different ways: (1) based on the obtained specific egg production rates, (2) a temperature dependent equation, and (3) a multilinear regression taking temperature, body weight and chlorophyll into consideration. The contribution of the small species was insignificant when compared to the large Calanus species during the spring- and post-bloom. However, during late summer and winter, where Calanus had left the upper water strata for hibernation, the small species played a crucial role in the pelagic carbon cycling.     

Euglenoid blooms in the floodplain wetlands of Barak Valley, Assam, North eastern India

Red blooms of Euglena sp. in the floodplain wetland ecosystems of Barak Valley, Assam, India, were found to be induced by high concentrations of NH3-N, NO3, Fe, Mg and to some extent, PO4, Cu and Zn in their water. The trace elements were rapidly accumulated by the bloom organisms to high levels, whereby their concentrations in the water declined, leading to a collapse of the bloom, which tended to reappear as decomposition again led to the release of the nutrients. The bloom also harboured fairly high density of certain other algae and zooplankton, thereby acting as a sub-system within the wetland ecosystem. The bloom is non-toxic and is exploited as a fish food by the fish-farmers who artificially induce a bloom for augmenting the growth of surface-feeding species of fishes.     

Community structure of the rocky shore macrobenthos in southern Hokkaido,Japan

Relationships between community structure and environmental factors were sought through multivariate analysis (quantification analysis and correspondence analysis) of data obtained in a benthic macrofauna survey of a rocky low-tide platform located at Usujiri, southern Hokkaido, Japan. A total of 21 species were found. However, their annual mean abundance was dominated by only a few species. Three dominant species of the platform (Chthamalus challengeri, Littorina brevicula andSeptifer virgatus) accounted for an average of 92% of total abundance. Three environmental factors with associated species groups were examined. Among them, micro-topographic characteristics and wave action defined the main correspondence variables.L. brevicula was a characteristic member of the boulder and sheltered field.S. virgatus was the dominant species of sloped surfaces.C. challengeri, S. virgatus, andMytilus edulis were characteristic of ledge or bench environments. Finally,C. challengeri was characteristic of nip and high wave-exposure environments, given its exclusive appearences in these locations. However, as the degree of exposure depends on the scale of wash through the platform, it is directly limited by the microtopographic properties of low-tide platforms. Specific environmental influences such as exposure and height above the datum-plane were rather disapparent. Any overall effects on the community structure were synthesized by those of microtopography. After complete consideration, it was concluded that the community structure of the macrobenthos in this low-tide platform was found to be primarily under the influence of microtopographic characteristics.     

Physico-chemical characteristics of the peloid mud from Morinje Bay (eastern Adriatic coast, Croatia): suitability for use in balneotherapy

The study conducted in the Morinje Bay (Central Dalmatia, Croatia) included the investigation of the sedimentological, physico-chemical and geochemical properties of the Morinje Bay sediments in order to assess the suitability of the material for wellness-related purposes including medicinal use. The sedimentological characteristics of the Morinje mud are in accordance with the geological origin of the material; the sediment is fine grained, composed mostly of carbonates, quartz and clay minerals. The composition of the sediment determines the physico-chemical properties of the mud. The high content of carbonate minerals diminishes adsorbing the characteristics of the mud to some extent; however, they are still comparable to some commercial mud already successfully used for medicinal purposes. Geochemical analyses have shown that trace metals concentrations in the Morinje Bay samples were mostly in the range with similar material from unpolluted Adriatic environments. Only the concentrations of Cu, Pb, Zn and As were slightly higher than in the reference material, but still within acceptable limits.     

Diel variation and seasonal consistency in the fish assemblage of the non-estuarine Sulaibikhat Bay,Kuwait

In the period between September 1986 and August 1988, using an otter trawl, a total of 50 species from 30 families were captured in Sulaibikhat Bay, Kuwait. The dominant species were Liza carinata (Valenciennes), Pomadasys stridens (Forrskal) and Leiognathus brevirostris (Valenciennes). Although the total number of fishes captured in 1986–1987 and 1987–1988 differed by an order of magnitude, the seasonal changes in numbers, biomass and number of species in the intertidal and subtidal zones were consistent. Unbalanced three-way analysis of variance showed that significantly larger numbers of fishes and number of species, but not biomass, were captured in the intertidal at night compared to the intertidal during the day. This difference may be due to both increased net efficiency and movement of fishes into the intertidal zone at night to avoid piscivorous fishes that move from deep water into shallower water. During the day few fish are present in the intertidal zone and this may be to avoid predation by piscivorous birds. Several species of fishes were shown to have different patterns of behaviour during the diel period.     

Comparative life histories of three species of planktonic calanoid copepods in Dabob Bay,Washington

The life history patterns of Calanus marshallae Frost, C. pacificus Brodsky, and Metridia lucens Boeck were determined in Dabob Bay, Washington, USA, during 1973 and October 1981–September 1982. C. marshallae emerged from diapause and moulted to adults in January–February. One major generation was observed, having been produced mainly during early to mid-March. Most surviving individuals spawned in March, arrested development at the C5 stage, and were in diapause by late May. However, some individuals developed to adults by late April and produced a very minor second generation. C. pacificus emerged from diapause and moulted to adults in February–March. The first generation produced during the spring lagged slightly behind the C. marshallae generation. C. pacificus produced two additional generations, one in late spring and one in the fall. The production of the three generations coincided with or closely followed phytoplankton blooms. Some C. pacificus arrested development at the C5 stage and entered diapause during the summer, but the majority of the population did not do so until fall. M. lucens did not appear to enter a diapause state. The fall and winter population was chiefly adult females which mostly remained at depth and were reproductively immature. Nevertheless, naupliar M. lucens were found on all sampling dates, indicating that reproduction never ceased as it did for the two Calanus species. M. lucens produced generations in late winter/early spring, late spring, and late summer/early fall. Interspecific comparisons between the life histories observed in Dabob Bay and intraspecific comparisons between the life histories in Dabob Bay and other locations are made. The advantages or disadvantages conferred to the copepods by their differing life histories are also discussed.