Reproductive behavior of the Atlantic silverside,Menidia menidia (Pisces,Atherinidae)

Synopsis The spawning behavior of the Atlantic silverside,Menidia menidia, was studied at two sites on the North Edisto River estuary in South Carolina. Prespawning schools moved back and forth along the shoreline as the time of high tide approached. Spawning runs took place in the upper intertidal zone at high tide.Spawning fish deposited their eggs on three types of substrates: 1) the lower stems of cordgrass plants,Spartina alterniflora, 2) detrital mats, and 3) exposed cordgrass roots along erosional scraps. Spawning behavior during egg deposition and fertilization was similar for all three substrates. Females released eggs during a rapid fluttering motion of the posterior half of the body. A similar movement accompanied release of sperm by males. Behavior of fish just prior to spawning insured deposition of gametes at locations that provided protection from thermal and drying stress during development. Eggs were deposited at mean intertidal elevations of 1.8 and 1.5 meters above mean low water (MLW) at respective study sites. They were exposed to the atmosphere for approximately ten hours between successive high tides.During spawning runs in which eggs were deposited at the base of cordgrass plants, ambient dissolved oxygen concentrations of the water in the spawning area were sometimes reduced to < 1.0 mg. 1^–1. Spent fish, apparently incurring an oxygen debt while spawning, formed a nonschooling aggregation offshore from the spawning zone.The recurrent use of specific spawning substrates for egg deposition resulted in an uneven distribution of spawning runs along the shoreline at each study site.Contribution No. 409 to the Gulf Breeze Environmental Research Laboratory. Contribution No. 377 to the Belle W. Baruch Institute for Marine Biology and Coastal Research.     

Spawning ecology of flannelmouth sucker, Catostomus lattipinnis (Catostomidae), in two small tributaries of the lower Colorado River

We report the first published accounts of spawning behavior and spawning site selection of the flannelmouth sucker in two small tributaries of the lower Colorado River in the Grand Canyon, Arizona. Spawning was observed on 20 March 1992 and from 28 March to 10 April 1993 in the Paria River, and from 16 to 19 March 1993 in Bright Angel Creek. Flannelmouth suckers exhibited promiscuous spawning behavior–individual females were typically paired with two or more males for a given event and sometimes changed partners between events. Multiple egg deposits by different females sometimes occurred at one spawning site. Flannelmouth sucker selected substrates from 16 to 32 mm diameter in both streams. Spawning occurred at depths of 10 to 25 cm in the Paria River and 19 to 41 cm in Bright Angel Creek. Mean column water velocities at spawning locations ranged from 0.15 to 1.0 m sec^-1 in the Paria River and from 0.23 to 0.89 m sec^-1 in Bright Angel Creek. Water temperatures recorded during spawning ranged from 9 to 18° C in the Paria River and 13 to 15° C in Bright Angel Creek. Spawning flannelmouth sucker ascended 9.8 km upstream in the Paria River and 1.25 km in Bright Angel Creek. Spawning females (410–580 mm) were significantly larger than spawning males (385–530 mm) in the Paria River. The mean size of spawning fish in the Paria River was significantly smaller than the entire stock, averaged throughout the study period (380–620 mm). However, fish spawning in 1992–1993 averaged 53 mm larger than fish spawning in the same reach of the Paria River in 1981, indicating a shift in the size structure of this stock.     

Environmental Biology of an Endemic Cyprinid,Varicorhinus alticorpus,in a Subtropical Mountain Stream of Taiwan

An endemic freshwater fish, Varicorhinus alticorpus, was studied from 1990 to 1993 in a fast flowing mountain stream in southern Taiwan (22°30'N, 120°30'E). The analysis of environmental conditions suggested that the fish could tolerate water temperatures between 19–24°C, a pH of 8.0–8.8, and flow velocity from 20–100cmsec^–1. The fish occupied riffles and pools. They foraged and schooled during daytime and aggregated in crevices between rocks at night. Most juveniles stayed at the shallow sandy-pebble flat by the pool where flow was slow. The species feed primarily on the periphyton growing on the rocks, and leave unique scars after grazing. One peak of juvenile recruitment was observed right after the rainy season, suggesting that reproduction of the fish has adapted to the climate. The change of abundance of adults along the habitats (at 150 to 800 m altitude) also indicated that the fish might migrate to low altitudes for spawning and disperse back to higher altitudes for exploiting new resources. The environmental biology of this fish shows an example of a cyprinid adapted to habitats in a subtropical mountain stream.     

Reproduction of the Nassau grouper,Epinephelus striatus (Pisces: Serranidae) and its relationship to environmental conditions

Synopsis Spawning aggregations of the Nassau grouper, Epinephelus striatus, occurring at the southern end of Long Island were investigated during the winters of 1987–1988 and 1988–1989. Fish aggregate at two specific sites around the time of the full moon in December and January, possibly not during November and almost certainly not during February. At one site the fish aggregate well inside the shelf edge while at the other they are on the dropoff into deep water. Fish migrate, probably as distinct groups of up to about 500 individuals, to the sites on or before the full moon and remain for several days. There was no significant difference in the size of males and females caught from the aggregations between years or months. Female/male sex ratio was between 5 : 1 and 3 : 1. Courtship occurred in late afternoon with spawning commencing shortly before sunset. Most spawning occurred within 10 min of sunset. Water temperatures were 25.0–25.5°C during a period of gradual decrease towards the annual minimum in February and March. Two color patterns were important in courtship and spawning. The bicolor pattern is a submissive coloration indicating a non-aggressive state acquired by both males and females near the time of spawning. The dark phase is acquired by females who are followed by numerous bicolor fish during courtship and they lead spawning events in this pattern. Spawning occurred among subgroups of the aggregation numbering 3–25 fish; gamete release was well above the bottom. Drogues deployed with the gametes either moved inshore or did not move far away from the shelf edge over the course of several days. Nassau groupers may not strictly be protogynous hermaphrodites and other groupers which form spawning aggregations may not be so either. Latitudinal shifts in spawning time may be related to water temperatures. Currents at aggregation sites do not appear to favor offshore transport of eggs. Questions of whether spawning aggregations should be protected need to be answered.     

Periodic movement,recruitment and size-related distribution of the sea cucumber Holothuria scabra in Solomon Islands

Field studies of the sea cucumber Holothuria scabra conducted in Kogu Veke, Solomon Islands, showed monthly recruitment of newly-settled juveniles on seagrass and indicated that size distribution was a function of substratum type and depth. Adults >250 mm body length were found mainly on sand, with <5% organic matter (OM), at depths of >1–3 m. Individuals >10–250 mm were found mostly in 30–120 cm of water, on mud and muddy sand with OM content between 5 and 10%. Specimens >40–150 mm were also found in the intertidal zone, sometimes burrowed on exposed sandflats at low tide. Holothuria scabra avoided substrata of fine silt or shell and coral pebbles, and sediment with an organic content 30%. Juveniles 100 mm burrowed at sunrise and surfaced at sunset, whereas individuals >100 mm burrowed and surfaced a few hours earlier. Holothuria scabra tended to burrow when salinity decreased, whereas increased water temperatures reduced normal burrowing behaviour. Spatial distribution, observed during tank experiments, suggested that adult H. scabra aggregated prior to spawning and in response to the lunar cycle. The formation of pairs, trios or larger groups increased during the new moon and was most common just before the full moon. Newly-settled juveniles up to ca. 9 mm were found on seagrass leaves. Typically, maximum densities and smallest recruits were observed a couple of weeks after the full moon, lower densities and slightly larger recruits were found a few days later. Juveniles with a mean length around 65 mm released on sand moved less and grew faster than juveniles released in seagrass beds or on substrata of shells and crushed coral.     

Diversity and cold-active hydrolytic enzymes of culturable bacteria associated with Arctic sea ice,Spitzbergen

The diversity of culturable bacteria associated with sea ice from four permanently cold fjords of Spitzbergen, Arctic Ocean, was investigated. A total of 116 psychrophilic and psychrotolerant strains were isolated under aerobic conditions at 4°C. The isolates were grouped using amplified rDNA restriction analysis fingerprinting and identified by partial sequencing of 16S rRNA gene. The bacterial isolates fell in five phylogenetic groups: subclasses and of Proteobacteria, the Bacillus–Clostridium group, the order Actinomycetales, and the Cytophaga–Flexibacter–Bacteroides (CFB) phylum. Over 70% of the isolates were affiliated with the Proteobacteria subclass. Based on phylogenetic analysis (<98% sequence similarity), over 40% of Arctic isolates represent potentially novel species or genera. Most of the isolates were psychrotolerant and grew optimally between 20 and 25°C. Only a few strains were psychrophilic, with an optimal growth at 10–15°C. The majority of the bacterial strains were able to secrete a broad range of cold-active hydrolytic enzymes into the medium at a cultivation temperature of 4°C. The isolates that are able to degrade proteins (skim milk, casein), lipids (olive oil), and polysaccharides (starch, pectin) account for, respectively, 56, 31, and 21% of sea-ice and seawater strains. The temperature dependences for enzyme production during growth and enzymatic activity were determined for two selected enzymes, -amylase and -galactosidase. Interestingly, high levels of enzyme productions were measured at growth temperatures between 4 and 10°C, and almost no production was detected at higher temperatures (20–30°C). Catalytic activity was detected even below the freezing point of water (at –5°C), demonstrating the unique properties of these enzymes.     

Tidal Influence on Spatial Dynamics of Leopard Sharks, Triakis semifasciata, in Tomales Bay, California

We used ultrasonic telemetry to determine the movement directions and movement rates of leopard sharks, Triakis semifasciata, in Tomales Bay, California. To analyze tide and time of day effects, we surgically implanted transmitters in the peritoneal cavities of one male and five female leopard sharks, which we located during summer for three to five sampling sessions lasting 12 to 24h each. All leopard sharks showed strong movement direction patterns with tide. During incoming tides, sharks moved significantly (p<0.0001) towards the inner bay, apparently to exploit the extensive inner bay muddy littoral zones' food resources. On outgoing tides, sharks showed significant (p<0.0001) movements towards the outer bay. During high tide, there was no discernible pattern to their movements (p=0.092). Shark movement rates were significantly (p<0.0001) greater during dark periods (mean±SE: 10.5±1.0m min^–1), compared with fully lighted ones (6.7±0.5m min^–1). Movement rates of longer sharks tended to be greater than those of shorter ones (range means±SE: 5.8±0.6m min^–1 for the 91cm shark, to 12.8±1.6m min^–1 for the 119cm shark), but the leopard sharks' overall mean movement rate (8.1±0.5m min^–1) was slower than other (more pelagic) sharks.     

Mean rate of DNA replication and replicon size in the shoot apex ofSilene coelirosa L. During the initial 120 minutes of the first day of floral induction

Summary 28-day-old plants ofSilene coeli-rosa were exposed, at 1,700 hours, to long day (LD) conditions comprising light of low fluence rate provided by tungsten bulbs, or maintained in darkness as short day (SD) controls. All plants were exposed at 1,700 hours to tritiated-(methyl-³H)-thymidine for 30, 45, 60, 90, or 120 minutes. Apical domes were isolated and prepared as fiber autoradiographs from which replicon size and rates of DNA replication, per single replication fork were recorded. In SD, replicon size was between 15–20 m and exposure to LD conditions altered neither replicon size nor the pattern of deployment of replicons during S-phase relative to the SD controls. However, the mean rate of replication in LD was 8.7 m h^–1 compared with 5.2 m h^–1 in SD. Thus, exposure to LD resulted in a 1.7-fold increase in the rate of DNA replication relative to the SD controls. This rapid increase in replication rate, detectable within 30 minutes of the start of the LD is discussed in relation to changes known to occur to the cell cycle inSilene during the first day of floral induction.     

Site fidelity, home range, and tidal migrations of juvenile pinfish, Lagodon rhomboides, in salt marsh creeks

We conducted a series of experiments to determine the movements and fidelity of juvenile pinfish, Lagodon rhomboides, in salt marsh creeks within North Inlet Estuary, SC. In Experiment 1, we investigated the fidelity of pinfish (40–100mm SL) at four subtidal locations (9–617m apart) along the axis of a major creek. We trapped and marked 2,297 individuals and recaptured 15–22% of the fishes released at each site, almost all of which (99.7%) had been marked at the same location of recapture up to 3 months earlier. In Experiment 2, we investigated pinfish movements between subtidal and intertidal areas. In the 8 weeks prior to sampling 2 intertidal creeks, we marked 950 juveniles in the adjacent subtidal areas. Sampling of the 2 flooded intertidal creeks showed that 9–20% of the pinfish collected bore marks, and all had been previously marked at the subtidal site immediately adjacent to the intertidal creek. Gut analysis of 60 individuals revealed that juveniles collected from the intertidal areas at high tide had consumed about ten times more food than those collected at the subtidal sites at low tide. In Experiment 3, we determined the fidelity of recently settled pinfish (<35mm SL) and showed that 15% of the 434 marked pinfish remained in the same area; some were at liberty up to 7 weeks. In Experiment 4, we determined home range by tagging and recapturing juvenile pinfish at sites separated by 20m along a 200m subtidal transect. We estimated that during periods when the intertidal zone was not accessible, the average home range was 9.4m with only 10% of tagged fish moving greater than 20m during the 4 month study. Our results indicate that soon after recruitment to the estuary, pinfish establish strong fidelity for sites within salt marsh creeks and exhibit tidal periodicity in both movements and feeding.     

Observations on the distribution patterns,behaviour, diets and reproductive cycles of sand-dwelling clinids (Perciformes: Clinidae) from South Africa

Synopsis Twenty-two samples of sand-dwelling fishes were collected from sublittoral sandy substrata off the Cape Peninsula, South Africa using a novel quantitative technique in which the ichthyocide, rotenone, was introduced beneath 6.25 m² plastic sheets weighted around the perimeter with chain. A total of 94 fish of eight species and four families were recorded. Of these, four species of the family Clinidae made up 92% of the material. Fish densities varied from 0.10–2.96 fish m^–2 (0.17–1.14 g m^–2). Distribution patterns amongst the Clinidae were explained by sediment particle size, with all of the newly discovered species, Cancelloxus longior, inhabiting fine sand (median grain size 0.25–0.50 mm), and all Xenopoclinus leprosus, C. elongatus and Pavoclinus smalei occurring in gravel (> 1 mm). Xenopoclinus kochi was found in all sediment types, but was most abundant in coarse substrata. Diets of all four species were similar, consisting chiefly of amphipods and isopods. However, small differences in prey preference were evident. All four species reached sexual maturity at 25–30 mm standard length, were viviparous, exhibited superembryonation and gave birth to live young of approximately 14 mm. Breeding seasonality occurred in all species except P. smalei.     

Mound-building and Feeding Behavior of the Twostripe Goby, Valenciennea helsdingenii, in the South Red Sea

The twostripe goby, Valenciennea helsdingenii, lives in pairs that cooperate in building large mounds, up to 60cm diameter, over their burrows. In 10-min observations (N=81), their activities were recorded by SCUBA divers (directly or on video) from 17 April to 6 May 1994 in the Jabal Zuqar and Hanish Islands, Yemen, in the south Red Sea. A pair of gobies moved up to 90 pieces of rubble in 10min (0–45, 8.4 pieces per fish per 10-min observation). Pairs generally stayed close together and alternated in feeding, one watching while the other grabbed a mouthful of sand most of which filtered out the gill openings. A pair fed up to 259 times in one 10-min observation period with an average rate of 3.7 (range 0–13) times per fish per minute. The feeding rate was lower at dusk when the pair's main activity was carrying sand and rubble that closed the entrance of their burrow before plunging through this soft new barrier to retire inside for the night. They emerged from their burrow about 40min after sunrise. Half of one medium-size mound was taken apart and consisted of 4062 pieces of coral rubble and shells. The hypothesis that building large mounds may be related to dominance and competition for mates, as in some cichlid fishes, does not apply to the isolated, stable pairs of V. helsdingenii.     

Morphology and infraciliature of Holosticha bradburyae nov. spec. (Ciliophora,Hypotrichida) from the Yellow Sea,China

The living morphology and infraciliature of a new marine hypotrichous ciliate, Holosticha bradburyae nov. spec., collected from the coastal water off Qingdao (Tsingtao), China, are investigated. This species is characterized by: body size 150–320 × 25–75 m with brownish to dark brown cell colour, ca. 53 adoral membranelles and 1 anteriorly positioned buccal cirrus; 3 frontal, 2 frontoterminal and 20–26 transverse cirri; midventral rows comprising 27-32 pairs of cirri; one conspicuous gap always present between anterior and posterior parts of AZM, and 2–5 distinctly elongated membranelles are always present at the posteriormost end; cortical granules conspicuous, round and flattened with central depression, arranged in about 10 lines on dorsal side; 28–33 irregularly arranged macronuclear nodules; 9–11 complete dorsal kineties; caudal cirri absent.     

Thresholds of retinal and extraretinal photoreceptors measured by photobehavioral response in catfish,Silurus asotus

Summary Light thresholds of retinal and extraretinal photoreceptors in catfish were examined by the photobehavioral response using a method in which reflex body movements are recorded. Thresholds were determined in six groups, (A) intact, (B) ophthalmectomized, (C) pinealectomized, (D) ophthalmectomized + pinealectomized, (E) ophthalmectomized, pinealectomized and skinless over the brain (skinless fish) and (F) ophthalmectomized, pinealectomized and dorsally covered with aluminum foil over the brain (covered fish). All these fishes displayed short term activity to white light stimulation after being dark adapted for more than 5 h. The lowest threshold was obtained in the intact group (2.0×10^–4 W/cm²). The thresholds of lateral eyes and the pineal organ were 3.4×10^–3 and 1.5×10^–2 W/cm², respectively. Without lateral eyes and pineal organ, catfish still responded to light, indicating the possible existence of extraretinal nonpineal photoreceptors (ENPs). The threshold of ENPs was 3.3 W/cm². The localization of ENPs was assumed to be in the brain from the experiment with the combination of skinless and covered fish.Abbreviation ENP extraretinal nonpineal photoreceptor     

Social system of an inshore stock of the red hind grouper,Epinephelus guttatus (Pisces: Serranidae)

Synopsis According to sex allocation theory, the decision by a female in protogynous fish species to change sex or not should be influenced by, among other things, the mating sex ratio during spawning periods and/or by factors that vary directly with the spawning sex ratio, such as relative rates of behavioral interaction with males and females outside of spawning periods. In groupers that only spawn during a few weeks of the year in large aggregations, individuals must assess the relative value of changing sex or not entirely within the aggregation unless the social system during the remainder of the year provides a behavioral equivalent of the mating sex ratio. Fifty-five individuals of the red hind,Epinephelus guttatus, were tagged and repeatedly located during a 152-day period within a 100 × 100 m grid on a shallow forereef off southwestern Puerto Rico. The home ranges of 22 tagged individuals sighted 10 or more times were 112–5636 m² in area. Individual home ranges overlapped with the home ranges of 1–18 other individuals. Home ranges of small fish were not clustered within the borders of the home range of larger fish, i.e. fish did not form spatially defined social units. At the end of the study, 31 tagged individuals remained on the grid together with five newly sighted fish. All 36 individuals proved on histological examination to be females similar in size to females in the spawning aggregation of the following year. The sex ratio of this all-female inshore stock differed significantly from the sex ratio of that spawning aggregation. Hence, information predicting the reproductive value of a sex change is not available to females in the inshore stock during nonspawning months.     

Growth of dark chub, Zacco temmincki (Cyprinidae), with a discussion of sexual size differences

I examined the annual and seasonal growth of dark chub, Zacco temmincki, in a Japanese river. Investigation of opercular rings showed that the fish reached a maximum age of 8 years. There was no significant sexual size difference at younger ages (3–5 years), but males were larger than females at older ages (6–7 years). Annual increments of length and weight for males that were recaptured were also larger than those for females. The fish spawn from June to August. Females grew for a short period from April to May, but males on average grew for a longer period from April to August. There was no sexual difference in growth rate except during the spawning period. Annual growth rate was negatively correlated with fish length in each sex. The sexual size differences at older ages of the fish might be due to the polygynous mating system in which most mature males could not obtain females and invested for somatic growth in the spawning period, and a short growing season that was overlapped considerably with the spawning period.     

Reproductive Biology of the Coral-reef Goby, Asterropteryx Semipunctata, in Kaneohe Bay, Hawaii

A population of the Hawaiian coral-reef goby, Asterropteryx semipunctata, was sampled over a 12-month period to determine basic demographics and reproductive parameters. The sexes did not differ in median length or weight, although the largest males were considerably longer and heavier than the largest females. Overall adult sex ratio was 1:1; monthly sex ratios did not differ from unity except in June, when there was a significant female bias. Minimum age at maturity (17.5–19mm SL) was estimated to be 4.5–5 months after hatching. Nearly all fish over 22mm SL were mature. Mature females were found in all months of the year, and females that showed evidence of recent or imminent spawning were collected in every month except December. Gonadal analyses indicated a peak in breeding during the summer (May–July) and minimal spawning during the winter (January–February). Between 20% and 30% of females showed evidence of having spawned within the 24-h period prior to collection; therefore, it was estimated that females spawned, on average, at least once every five days and perhaps as frequently as every three days. Mean batch fecundity was 708 eggs (± 418), and was not well predicted by standard length, body weight, or somatic condition. Relative batch fecundity (mean = 1.44 oocytes mg^–1 somatic wet weight) varied seasonally, with higher values in spring and summer (April–July) than in fall and winter (September–January). Reproductive parameters are compared and contrasted with those of other gobiid fishes to elucidate general differences between temperate and tropical species.     

Substrate preference in age-0 red snapper, Lutjanus campechanus

We tested age-0 red snapper, Lutjanus campechanus, for sand or shell substrate preference in a circular tank (1.5 m diameter × 0.6 m deep). The test tank was divided into two equal areas of whole oyster shell or sand substrates. All trials were video taped for 20 to 25 min. Tapes were viewed on a monitor and locations of all fish recorded and timed with respect to substrate. Mean ± SE time on shell was 11.6 ± 0.5 sec (4 fish trial^–1) and 13.8 ± 1.1 sec (1 fish trial^–1). Mean ± SE time on sand was 9.7 ± 0.4 sec (4 fish trial^–1) and 8.9 ± 0.5 sec (1 fish trial^–1). Fish spent significantly more time on shell compared to sand substrate in four-fish trials (paired t-test, p < 0.10) and also in single-fish trials (paired t-test, p < 0.05). Shell substrate may offer increased food and shelter for age-0 red snapper compared to sand substrate. However, a preference for shell substrate in the natural habitat may occur only during the nursery period, because as fish grow (> 100 mm TL) in the late fall they probably outgrow the shell habitat.     

Larval development of <Emphasis Type="Italic">Achoerodus viridis</Emphasis> (Pisces: Labridae), the Australian eastern blue groper

Larvae of the southeastern Australian endemic hypsigenyin labrid Achoerodus viridis are described and illustrated from 30 specimens (3.0–9.6mm) captured in plankton nets and larval fish beach seines. Development is typical for labrids. Larvae of A. viridis can be distinguished from those of other labrids because they possess distinctive pigment, 28 myomeres, and fin ray counts of D XI,11, A III,11, and P₁ 16–18. Larvae of closely related hypsigenyin labrid genera are poorly known but are similar in morphology and pigmentation. Adult A. viridis live on coastal rocky reefs. Larvae of 3.0–8.2mm were captured over the continental shelf off central New South Wales in the Western Tasman Sea. Larvae of 6.6–7.7mm were captured in a tidal channel leading to an estuarine lagoon, and the smallest larvae captured in seagrass beds in the lagoon were 7.2 and 8.5mm.     

Denitrification in a seashore sandy deposit influenced by groundwater discharge

The chemical compositions of ground water and organic matter in sediments were investigated at a sandy shore of Tokyo Bay, Japan to determine the fate of ground water NO₃ ^–. On the basis of Cl^– distribution in ground water, the beach was classified into freshwater (FR)-, transition (TR)-, and seawater (SW)-zones from the land toward the shoreline. The NO₃ ^– and N₂O did not behave conservatively with respect to Cl^– during subsurface mixing of freshwater and seawater, suggesting NO₃ ^– consumption and N₂O production in the TR-zone. Absence of beach vegetation indicated that NO₃ ^– assimilation by higher plants was not as important as NO₃ ^– sink. Low NH₄ ⁺ concentrations in ground water revealed little reduction of NO₃ ^– to NH₄ ⁺. These facts implied that microbial denitrification and assimilation were the likely sinks for ground water NO₃ ^–. The potential activity and number of denitrifiers in water-saturated sediment were highest in the low-chlorinity part of the TR-zone. The location of the highest potential denitrification activity (DN-zone) overlapped with that of the highest NO₃ ^– concentration. The C/N ratio and carbon isotope ratio (¹³C) of organic matter in sediment (< 100 -m) varied from 12.0 to 22.5 and from –22.5 to –25.5, respectively. The ¹³C value was inversely related to the C/N ratio (r ² = 0.968, n = 11), which was explained by the mixing of organic matters of terrestrial and marine origins. In the DN-zone, the fine sediments were rich in organic matters with high C/N ratios and low ¹³C values, implying that dissolved organic matters of terrestrial origin might have been immobilized under slightly saline conditions. A concurrent supply of NO₃ ^– and organic matter to the TR-zone by ground water discharge probably generates favorable conditions for denitrifiers. Ground water NO₃ ^– discharged to the beach is thus partially denitrified and fixed as microbial biomass before it enters the sea. Further studies are necessary to determine the relative contribution of these processes for NO₃ ^– removal.