Seasonal fluctuations in the abundance of populations of marine cladocerans and their resting eggs in the Inland Sea of Japan

Seasonal fluctuations in the abundance of resting eggs of four species of marine cladocerans, Penilia avirostris Dana, Podon polyphemoides (Leuckart), Evadne tergestina Claus and Evadne nordmanni Lovén, were investigated in association with their planktonic populations in the central part of the Inland Sea of Japan for a period of over two years. Most numerous were the eggs of P. avirostris, followed by those of the species in the aforementioned order. There was a marked seasonal fluctuation in the number of eggs that was closely correlated with the occurrence of the planktonic population for each species. The egg-number was highest just before the planktonic population disappeared from the water column, and gradually decreased thereafter until a minimum value was reached shortly before the new planktonic population appeared in the next season. The pattern of fluctuation in number was found to be very similar among all four species, indicating strongly that the cladoceran populations in this warm temperate sea may persist either as plankton in warmer seasons or as benthic resting eggs in colder seasons. The possible causes of the decrease in egg density in bottom sediments are discussed.     

Feeding habits of marine cladocerans in the Inland Sea of Japan

Natural food items of five species of marine cladocerans, Evadne nordmanni, E. tergestina, Penilia avirostris, Podon leuckarti and P. polyphemoides, were investigated in the Inland Sea of Japan between April 1986 and May 1987. Gut content examination with SEM (scanning electron microscopy) revealed that feeding was largely limited to centric diatoms and a few exceptions of pennate diatoms and dinoflagellates. No animal remains were detected, and some unidentified materials were also found. Phytoplankton smaller than 35 m in size (cell diameter in centric diatoms and longest dimension in others) was found most frequently in the gut of cladocerans. The role of grazing of marine cladocerans in trophodynamic pathways of the pelagic realm is discussed.     

Distribution and abundance of ophiuroids on the continental shelf and slope of the East Sea (southwestern Sea of Japan), Korea

The present study was carried out to describe the distribution of ophiuroids on the continental shelf and slope of the East Sea, off the Korean Peninsula. Three cruises were made from 1985 to 1987 in the area between 35 to 38°N and 129 to 130°E. A total of ten species was identified from the samples collected at 136 stations. Among them four species (Amphiodia craterodometa, Amphioplus macraspis, Ophiura leptoctenia and Ophiura sarsi) constituted more than 98% of abundance by number. The continental shelf off Gampo, near 35°40N, could be considered to be the southern limit of cold-water species. The abundance of dominant ophiuroids changed according to depth. Three faunal zones, namely, the inner shelf, the shelf edge and the slope zones, coincided with the topographic divisions of continental shelf and slope. The major inhabitant of the inner shelf was A. craterodometa and that of the continental slope was O. leptoctenia. The shelf edge area revealed a transitional zone where all four species occurred, contributing to the total number of individuals with almost equal proportions. The temperature of bottom waters was considered to be a factor determining zonal distribution. The sediment characteristics were not closely related to ophiuroid distribution.     

Ontogenetic diel vertical migration of the planktonic copepodCalanus sinicus in the Inland Sea of Japan

The ontogenetic diel vertical migration of the planktonic copepodCalanus sinicus was investigated in the Inland Sea of Japan in summer 1988, when the water was thermally stratified with a thermocline of ca 5 °C between 35 and 45 m. Stage-specific differences in the diel vertical migration behavior ofC. sinicus were found. Eggs were spawned primarily within the surface-waters between midnight and dawn by ascending females, and sank gradually to deeper waters until they hatched into nauplii. Non-feeding nauplius stages (NI and II) were distributed throughout the water column, but the first feeding stage (NIII) performed an ontogenetic upward migration. NIV to VI and copepodite (C) stages I to III continuously aggregated in the phytoplankton-rich euphotic layer. However, the depth of the median CI to III populations descended as stage progressed. The onset of prominent diel vertical migration took place in CIV, and the amplitude of vertical migration increased with age, being maximal in adult females (CVI). Adult males (CVI), however, remained in the layer below 20 m, and did not migrate dielly. The ecological significance of ontogenetic diel vertical migration is discussed.     

Ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus in the Inland Sea of Japan

The ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus was investigated in the Inland Sea of Japan in June 1989, when the water column was thermally weakly stratified. Because of fewer eggs and less variation in their abundance, nocturnal spawning was not apparent. A pronounced upward migration occurred in NIII. NIII to CIII resided in the upper 20 m layer throughout the day, and from CIV on their median depths descended. CV and adult females underwent significant diel vertical migration, whereas adult males did not migrate. By integrating the results from the present study and those from our previous investigations (in August–September 1988, November 1988 and March 1989), we review seasonal variation in the ontogenetic diel vertical migration of C. sinicus. Spawning was largely nocturnal, reaching its maximum level around dawn, but spawning depth and fecundity changed seasonally. The distribution of pre-feeding stages, NI and NII, was similar to that of eggs. A pronounced upward migration always occurred in the first feeding stage, NIII, and late nauplii and early copepodites always resided in the food-rich upper layer, indicating that upward migration by NIII is feeding migration. As the stages progressed, they extended their vertical distribution range, and CV and adult females usually underwent diel vertical migration. However, the pattern and strength of this migration differed seasonally. Their day depths increased with the increase of relative biomass of planktivorous fish, indicating that predator avoidance induces their diurnal downward migration. High chlorophyll a concentrations in the upper layer (<15 m deep) relative to the lower layer (>20 m deep) amplified their diel vertical migrations. Diel vertical migration of C. sinicus is a phenotypic behavior.     

Ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus in the Inland Sea of Japan

The ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus was investigated in the Inland Sea of Japan in November 1988 and March 1989, when the water temperature was weakly stratified in a reversed manner. In both investigations a pronounced ontogenetic difference in vertical distribution was found. Spawning always occurred during nighttime, being confined to the upper 40 m water column in November but to the layer below 35 m in March. The distribution of pre-feeding nauplius stages, NI and NII, was more or less similar to that of the eggs. The first-feeding NIII performed a marked upward migration, and late nauplius stages (NIV to NVI) and early copepodite stages (CI and CII) continuously aggregated in the upper water column where phytoplankton was abundant. CIII to CVI (adult female and male) tended to disperse in the whole water column. In November, however, they avoided the upper 10 m strate during daytime and some individuals migrated upward to the surface during nighttime. In March, CV and CVI aggregated in the layer between 5 and 15 m deep in the daytime and migrated both upward and downward at dusk, resulting in homogeneous distributions during the nighttime.     

Calanoid copepod eggs in sea-bottom muds. II. Seasonal cycles of abundance in the populations of several species of copepods and their eggs in the Inland Sea of Japan

The seasonal cycles of abundance of populations of dominant calanoid copepods in the water column and of their eggs recovered from the bottom sediment in the central part of the Inland Sea of Japan are described. The numbers of both copepods and eggs fluctuated markedly with season in an essentially similar pattern among the 6 species studied (Tortanus forcipatus Giesbrecht, Calanopia thompsoni A. Scott, Acartia erythraea Giesbrecht, A. clausi Giesbrecht, Centropages abdominalis Sato, C. yamadai Mori). The density of eggs in the sea bottom was highest shortly before the population of adults and late copepodids disappeared from the plankton; the numbers of eggs then gradually decreased until the appearance of the next planktonic population.     

Growth and production of the inshore marine copepod Pseudodiaptomus marinus in the central part of the Inland Sea of Japan

The growth and production of the inshore marine copepod Pseudodiaptomus marinus was studied in the central part of the Inland Sea of Japan. The stage-specific growth rate was determined under controlled laboratory conditions by examining the length-weight relationship and development rates at various temperatures. The stage duration was short and constant from NII to CII, beyond which development was retarded. Males developed faster than females in CIV and CV. The specific growth rate was highest in copepodite stages followed by the nauplii and adult females (=egg production rate). The daily production of P. marinus was estimated from the stage-specific growth rate and stage-specific abundance in nature as the sum of the individual stages. The production changed seasonally with water temperature and population biomass. Daily production and biomass (P/B) ratios increased linearly with temperature. Total annual production was 20.7 mg C m^-3 yr^-1.     

Synchronous recruitment and growth pattern of planktonic larvae of the amphioxus Branchiostoma belcheri in the Seto Inland Sea, Japan

Planktonic larvae of the amphioxus Branchiostoma belcheri were collected in the western part of the Seto Inland Sea, Japan, in order to document their seasonal occurrence, recruitment, and growth patterns. The larvae appeared from mid-July to early October and their size ranged from 0.6 to 5.5 mm. Three distinct cohorts were observed during the summers, indicating multiple, synchronous spawning within the population. The length increment pattern of a cohort demonstrates slow growth for at least a week during the early larval stage. July 10, 20, and 30 are the inferred dates when spawnings started to produce the three cohorts in 2001. Favorable temperature range for spawning was 21–23°C. Bias in population structure due to advection can be assumed to be negligible; therefore the length increment of the cohort, about 0.05 mm day⁻¹, is regarded as the larval growth rate after the initial slow-growth stage. The planktonic stage of the first cohort in 2001 is estimated to have lasted around 55 days. The relative proportions of the second and third cohorts in samples from surface and mid-depth waters suggest downward migration of the planktonic larvae, as they age.     

Annual life cycle of Chattonella spp., causative flagellates of noxious red tides in the Inland Sea of Japan

Effects of storage temperature on the dormancy and maturation of dormant cells of Chattonella spp. were investigated by the extinction dilution method using sediment samples collected in 1984 after the blooming in Harima-Nada. Seasonality of germinability in fresh sediments was also studied in 1985. A storage period of more than 4 months with a low temperature (11°C) was essential for the maturation to induce the germination capacity of dormant cells. The optimal temperature for germination (22°C) had no effects on the maturation. Using fresh Suo-Nada sediments, a marked seasonality of germinability was confirmed. It was weak from autumn to early winter, then strengthened gradually up to a high level, which was maintained between spring and early summer, and again decreased rapidly during summer. The annual life cycle of Chattonella spp., including vetetative and dormant phases, was summarized as follows: (1) vegetative cells in early summer originate from germination of dormant cells in sediments; (2) they form overwintering dormant cells during the summer season; (3) dormant cells spend a period of spontaneous dormancy until next spring; (4) the duration of post dormancy, an enforced one due to low temperatures, follows until early summer. The life cycle of Chattonella spp. is therefore well adapted to the temperature regime in temperate seas such as the Seto Inland Sea of Japan.     

Seasonal reproductive biology of the egg-carrying calanoid copepod Pseudodiaptomus marinus in a eutrophic inlet of the Inland Sea of Japan

In situ egg production of the egg-carrying calanoid copepod Pseudodiaptomus marinus was investigated in Fukuyama Harbor, a eutrophic inlet of the Inland Sea of Japan, at 3- to 5-d intervals for a year. This species reproduced throughout the year, and the adults showed a large abundance peak in June/July and a small peak in September/October. Females usually outnumbered males, comprising 61.4% of the annual mean. The composition of ovigerous females varied from 7.9 to 100%, with an annual mean of 55.7%. Adult prosome length was consistently large throughout winter and spring, and decreased with increasing temperature in summer and fall. Egg diameter varied from 98 to 121 μm, and was negatively correlated to temperature. The seasonal variation in clutch size (range: 15.1 to 38.2 eggs) was bicyclical, with peaks in May and December. The egg production rate of breeding females was low in January to March (mean: 2.3 eggs female⁻¹ d⁻¹), while it was constantly high from mid-May to early October (mean: 12.1 eggs female⁻¹ d⁻¹). The specific egg production rate for the breeding females was highly correlated to temperature; it increased linearly from 0.03 d⁻¹ at 9 °C to 0.27 d⁻¹ at 26 °C. Compared to other co-occurring copepods, the reproductive rate of P. marinus was lowest, which is one of the reasons why this species never dominates in this inlet. Received: 11 November 1996 / Accepted: 7 December 1996     

Distribution and abundance of choanoflagellates (Acanthoecidae) across the ice-edge zone in the Weddell Sea,Antarctica

Choanoflagellates are thought to be an important component of oceanic microbial food webs, but little quantitative data exists on their abundance,, distribution, or relationship to potential food sources. In an Antarctic ice edge zone (northern Weddell Sea, March 1986), choanoflagellate abundance varied over two orders of magnitude in the upper 100 m. The lowest abundances were recorded at the bottom of the water column under ice cover and the highest abundances occurred in the upper 30 m of open water. Species that were predominantly in colonies dominated the open-water samples. Abundances of total choanoflagellates and some individual species were correlated with primary and secondary biomass and production, indicating a response to gradients in potential food sources. This suggests that choanoflagellates are tightly coupled with their food sources and supports the contention that they may an important link between bacteria-sized particles and metazoan grazers.     

Distribution and abundance of moon jellyfish (Aurelia aurita) and its zooplankton food in the Black Sea

The distribution of moon jellyfish (Aurelia aurita Linnaeus, 1758) in the Black Sea was determined from plankton samples collected above the anoxic zone (maximum depth 200 m) in the summer, winter and spring during 1991–1995. Distribution was patchy. Average biomass ranged from 98 to 380 g m⁻², and abundance varied from 2 to 14 individuals m⁻². Biomass and abundance peaked in late spring and summer. The distribution was correlated with hydrographic features in the Black Sea, with higher concentrations occurring at the peripheries of anticyclonic eddies. Centers of the two main cyclonic gyres generally had a low biomass of A. aurita. From July 1992 to March 1995, the populations were largely concentrated in offshore regions. A. aurita were confined to the upper part of the mixed layer. Smaller A. aurita (≤1 cm) were present in early spring (March), and individuals reached maximum size in the summer. Release of the epyhrae occurred in spring on the northwestern shelf of the sea when the seawater temperature was 11–12 °C. Microscopic analysis of stomach contents showed that copepods and mollusks form their main diet. Received: 3 September 2000 / Accepted: 29 September 2000     

Distribution and abundance of ctenophores and their zooplankton food in the Black Sea. II. Mnemiopsis leidyi

The distribution of Mnemiopsis leidyi Agassiz, 1865 in the Black Sea was determined using plankton samples collected above the anoxic zone (maximum depth 200 m) in the summer, winter, and spring from 1991 to 1995. Distribution was patchy. Average biomasses of 15 to 500 g m⁻² were measured, and abundances varied from 10 to 180 ind m⁻². Biomass and abundance peaked in winter, and there was a secondary peak in the summer. The distribution of M. leidyi was correlated with hydrographic features in the Black Sea with higher concentrations in anticyclonic gyres. The centers of the two main cyclonic gyres generally had a low biomass of M. leidyi. From July 1992 to March 1995, the populations were largely offshore. M. leidyi were confined to the upper part of the mixed layer both day and night. Some individuals displayed a negative taxis to daylight and were concentrated below the thermocline at night. Smaller M. leidyi (1.5 to 2 cm) were present in the winter, and individuals reached maximum size in the summer. Although reproduction was continuous throughout the year, there were two distinct peaks: the larger peak in the summer and the smaller peak in the winter. Microscopic analysis of stomach contents showed that copepods and molluscs form their main diet. Received: 1 November 1997 / Accepted: 30 August 1999     

Population dynamics and production of the planktonic copepods in a eutrophic inlet of the Inland Sea of Japan. II. Acartia omorii

Population dynamics and production of the calanoid copepod Acartia omorii Bradford were studied from November 1986 to November 1987 in Fukuyama Harbor, a eutrophic inlet of the Inland Sea of Japan. This species was present in the plankton from October to July (temperature range: 8.9 to 24.3°C), with peaks in February-March and June. During this period, nine generations could be detected, for which the mean population egg production rate and midstage abundance of each life stage older than naupliar Stage (N) II were determined to trace survival. The population suffered extremely high mortality during the early life stages: on average only 2.5% of the eggs produced recruited into NII. This large loss is probably concentrated within the egg stage, due to predation, including cannibalism, by omnivorous copepods, in addition to sinking loss of eggs in the water column. However, the mortality from NII to copepodite Stage (C) V was negligible, indicating low predation pressure by large carnivores. The biomass of A. omorii showed marked seasonal variations in parallel with numerical abundance. The instantaneous growth rate of each stage increased exponentially with increasing temperature. The integrated production rate of A. omorii from 7 November 1986 to 21 July 1987 was 749 mg Cm^-3 or 5.62 g Cm^-2     

Population dynamics and production of the planktonic copepods in a eutrophic inlet of the Inland Sea of Japan. I. Centropages abdominalis

Population dynamics and production of the calanoid copepod Centropages abdominalis were studied from November 1986 to November 1987 in Fukuyama Harbor, in the central part of the Inland Sea of Japan. This species was present in the plankton during a cold-water period from November to June (temperature range: 8.9 to 21.1 °C), with a peak abundance (23 600 ind m^–3) in February. During this period, six generations could be detected, and each generation time agreed well with that predicted from food-satiated laboratory experiments, indicating that the natural population was not food-limited. The population suffered extremely high mortality during the period from egg to naupliar stage (N) II: only 0.02 to 4% of the eggs survived to NII. However, the mortality in stages older than NII was almost negligible. The growth rate of C. abdominalis increased exponentially with increasing temperature. Its biomass and production rate showed marked seasonal variations largely in parallel with numerical abundance. The estimated production between 7 November 1986 and 29 May 1987 was 355 mg C m^–3 or 2.66 g C m^–2, 95% of which occurred during February and March. The daily production rate to biomass ratio increased exponentially with temperature from 0.18 at 8.9°C to 0.37 at 19°C.     

Distribution and abundance of ctenophores, and their zooplankton food in the Black Sea. I. Pleurobrachia pileus

The distribution of Pleurobrachia pileus Müller, 1776 in the Black Sea was determined using plankton samples collected above the anoxic zone (maximum of 200 m) in the winter, spring, and summer of 1991 to 1995. The summer samples were collected in 1991 to 1993 (for a previous) and are included in this paper for comparative purposes. High concentrations of P. pileus were found at the northern edges of anticyclonic eddies along the southern coastal regions. The biomass and abundance of P. pileus increased from winter through spring to a peak in summer. The highest mean wet weight during a sampling period was 250 g m⁻², while the maximum wet weight was 1429 g m⁻². P. pileus was mostly found in a layer extending from the lower parts of the thermocline down to the anoxic zone, where the temperature was <8 °C. The vertical distribution of P. pileus biomass had two clear maxima at night: an upper maximum at 20 to 40 m was less pronounced than the lower maximum at 90 to 120 m depth. Mean body length of P. pileus did not exceed 12 mm. Smaller individuals (9 to 10 mm length) occurred in winter. P. pileus had two length classes in early spring (March 1995) and late summer (August 1993), indicating the presence of both newly hatched and larger individuals. Overall, the stomach contents of P. pileus consisted mainly of Copepoda (90%), Cladocera (1%), Mollusca (1%), fish eggs and larvae (1%), and other taxa (7%). The preferred food of P. pileus (frequency of occurrence) was: Calanus euxinus (39%), Pseudocalanus elongatus (30%), Acartia clausi (28%), Oithona similis (2%), and Paracalanus parvus (1%). The endoparasite Hysterothylacium aduncum was commonly found in P. pileus. Abundances of Mnemiopsis leidyi and P. pileus were either negatively correlated (r = −0.5 to −0.7) or positively correlated at a low significance level (r = 0.25 to 0.3) with abundance of A. clausi in different months of the year. Aurelia aurita abundance was correlated mainly with the abundance of C. euxinus from June 1991 to March/April 1995. Over the same period the abundance of P. pileus was significantly correlated with the abundance of P. elongatus, an important prey species. Received: 1 November 1997 / Accepted: 30 August 1999     

Population dynamics and production of the planktonic copepods in a eutrophic inlet of the Inland Sea of Japan. III.Paracalanus sp.

Population dynamics and production of the calanoid copepodParacalanus sp. were studied from November 1986 to November 1987 in Fukuyama Harbor, a eutrophic inlet of the Inland Sea of Japan. This species was perennial, with a large abundance peak in June/July and small peaks in September/October and November/December. During a year of investigation, 15 generations Gould be detected. For each generation, the mean population egg production rate and the mean daily midstage abundance front NIII to CV were determined to obtain a survival curve from egg to CV. The mortality was extremely high during the early life stages: on average only 7.1% of the eggs produced might survive into NIII. This high mortality might be caused by predation by sympatric omnivorous copepods, in addition to sinking loss of eggs from the waten column. The biomass ofParacalanus sp. showed marked seasonal variations largely in parallel with numerical abundance. The instantaneous growth rate of each developmental stage increased exponentially with temperature up to 20 °C, above which the rate was constant. The annual integrated production rate was 734 mg C m^–3 yr^–1 or 5.5 g C m^–2 yr^–1.     

“Seed bank” of coastal planktonic diatoms in bottom sediments of Hiroshima Bay, Seto Inland Sea, Japan

Abundance and temporal distribution of viable (able to germinate) resting stage cells of planktonic diatoms in bottom sediments have been investigated almost monthly during 1989 to 1992 in Hiroshima Bay, western part of Seto Inland Sea, Japan. The abundance of viable resting stages in bottom sediments was enumerated with the extinction dilution method (most probable number method, MPN). In bottom sediments of Hiroshima Bay, dominantly distributed species and/ or genera of the diatom resting stages were Skeletonema costatum, Chaetoceros spp. and Thalassiosira spp. Viable resting stages of these diatoms were densely distributed on the orders of 10³ to 10⁶ (MPN g⁻¹ wet sediments), and persisted in bottom sediments throughout the investigation period. Conversely, vegetative cells of these diatoms fluctuated remarkably in the water column and disappeared sporadically. Survival of the resting stages in a collected sediment sample was also determined with the MPN method, at different storage temperatures (5, 10, 15, 20, 25 °C). The survival test demonstrated that the diatom resting stages could survive in the dark for several months or years in sediments. Resting stages survived longer at the lower storage temperature, and the order of longevity was consistent within three diatoms (Chaetoceros spp. > Thalassiosira spp. > S. costatum) at each storage temperature. The present study suggests that these diatom resting stages in the coastal bottom sediments could serve as a “seed bank”, analogous to those of terrestrial plants. The seed bank would ensure the survival of diatoms within highly fluctuating coastal environments, while it would also be the source of sporadic and autochthonous diatom blooms in coastal waters. Received: 29 November 1996 / Accepted: 16 December 1996