Larval and juvenile fishes occurring with flood tides on an intertidal mudflat in the Tama River estuary, central Japan

Intertidal movements of fish larvae and juveniles on a mudflat in the Tama River estuary, central Japan, were investigated by comparing the abundance and sizes of fishes caught in the intertidal zone during flood tides with those in the subtidal zone during low tides. A total of 28465 individuals, belonging to 9 families and 20 species, were collected by small purse seine. Among the abundant species, planktonic larvae and juveniles of gobiids and Konosirus punctatus were more abundant in the intertidal zone at flood tide than the subtidal zone at low tide. Similar occurrence patterns were found in juvenile Plecoglossus altivelis altivelis and Lateolabrax japonicus, having fully developed swimming abilities. In contrast to these species, much higher abundances of epibenthic juveniles of 2 gobiids (Acanthogobius flavimanus and Gymnogobius macrognathos) were found in the subtidal zone at low tide, although they also utilized the intertidal zone at flood tide.     

The climbing behaviour of Cerithidea anticipata (Mollusca: Gastropoda): The roles of physical and biological factors

Abstract Many molluscs in tidal wetlands climb trees as the tide rises, a behaviour usually assumed to be a means of avoiding subtidal predators. Some species are more active during neap tides, when the access of subtidal predators to the forest is limited, but rest on trees during spring tides. Cerithidea anticipata, which inhabits the mangrove forests around Darwin Harbour (Northern Territory, Australia), displayed almost exactly the opposite pattern. This species climbed higher, and was less active, during neap tides that did not flood the forest than during spring tides. In experiments with tethered snails, individuals prevented from climbing died during neap tides, apparently from physiological stress. Further, individuals resting on trees around clearings, usually selected shaded sites. These results suggest that the major reason C. anticipata climbed was to avoid physiological stress during neap tides, not subtidal predators during spring tides. There was some evidence of predation under the canopy, but the rate was relatively low and the species responsible appeared to be resident in the forest.     

Foraging strategy of the mudskipper Periophthalmus sobrinus Eggert in a Kenyan mangrove

The feeding behaviour of Periophthalmus sobrinus was studied in a population living along the channels of the Tana river delta. In November 1993 mudskippers were sampled every 2 tidal h for two consecutive spring and neap tides both during the day and night. The fishes were sexed and their stomach contents analysed. The results indicated that the mudskipper has a carnivorous diet and forages during both day and night although feeding greatly decreases after dark. Quantitative and qualitative differences in diet were found during the day and night of the two synodic and tidal phases. These differences were also demonstrable in the two sexes. The study showed that females of P. sobrinus are more selective than males and that differences between the two sexes may be related to reproductive condition. Furthermore, the mudskippers showed a tidal rhythm in feeding behaviour with peaks of activity around low tide (LT) on spring tides and around high tide (HT) on neap tides. Foraging activity seems to be influenced by the different spatial and temporal distributions of the mudskipper's prey caused by synodic, tidal and diel changes in the environment.     

Foraging activity of the sand beach isopod Scyphax ornatus Dana

Abstract

The terrestrial isopod Scyphax ornatus lives on exposed sand beaches. Adult isopods spent the daytime in burrows near the high water mark and made nightly foraging excursions over the uncovered middle beach. Hourly records of their night time distribution showed that Scyphax congregated near the water’s edge during the last 4 h of flood tide, where they fed on carrion. Insects (mostly drowned honey bees) were the main food item, but coelenterates, amphipods, crabs, and goose barnacles were also eaten. During ebb and early flood tides Scyphax occurred at low densities over the uncovered middle beach. Analysis of the distribution of large and small food items on the beach showed that by feeding during the flood tide, Scyphax exploited a rich concentration of carrion that accumulated in the swash zone and that was moved slowly up the beach by the incoming tide.     

Spring-neap cycle as a major driver of temporal variations in feeding of intertidal fishes: Evidence from the sea catfish Sciades herzbergii (Ariidae) of equatorial west Atlantic mangrove creeks

On macrotidal coasts, short- to medium term variations in feeding of intertidal fishes are influenced by several interacting time scales. To identify the driver of major variations in the feeding habits of intertidal fish, we used the pemecou sea catfish Sciades herzbergii (Ariidae), an abundant intertidal benthic second order consumer, as a model species. We analyzed the influence of the spring-neap and the day-night cycle on intertidal abundance, stomach fullness, diet composition and food consumption of S. herzbergii using block nets set at slack high tides in two mangrove creeks in north Brazil. At spring tides, intertidal abundance, stomach fullness, and total daily consumption of S. herzbergii were on average 8.8, 1.9, and 3.8 times higher than at neap tides, respectively. At spring tides, Uca spp. and Grapsidae (mostly Pachygrapsus gracilis) dominated the diet, irrespective of the time of day. Other important food items were Insecta and the semiterrestrial crab Ucides cordatus. At neap tides, Capitellidae contributed to the diet of nightly inundations while no specimens were caught at daytime. Creek location had no effect on any variable. Results from our study area and evidence from other studies suggest that the spring-neap tide pulse is likely the major driver of short- to medium term variations in feeding of intertidal fishes. This has important implications for feeding-related issues on macro- and probably also on mesotidal coasts: (i) juvenile fishes may have fortnightly growth spurts, (ii) intertidal prey populations may suffer regular fluctuations in mortality, (iii) studies of the feeding ecology of intertidal fishes should cover the combined effects of the tidal, diel, lunar and seasonal cycles, and (iv) the modeling of food webs should consider the differences between highly dynamic spring tide and quieter neap tide conditions.     

Activity patterns, feeding and burrowing behaviour of the crab Ucides cordatus (Ucididae) in a high intertidal mangrove forest in North Brazil

Activity patterns, feeding and burrowing behaviour of the economically important semi-terrestrial mangrove crab Ucides cordatus (Ucididae, L. 1763) was studied in a high intertidal Rhizophora mangle forest stand in Bragança, North Brazil. Video observations in the rainy and dry season were conducted over 24 h cycles at different lunar phases to investigate the behaviour of these litter-feeding crabs outside their burrows. During the rainy season, crabs stayed inside their burrows for 79% and 92% of the time during day and night, respectively. Time spent for feeding, burrowing and other activities outside their burrows was significantly longer during the day with 9.9% (night: 1.7%) and at waning and waxing moon with 9% (full and new moon: 0.9%). At neap tides (no tidal inundation) foraging and feeding activities outside burrows were clearly light-dependent, increasing at dawn and decreasing at dusk. Highest activities during daytime relate to the visual localisation of food. During the dry season, crabs spent less time inside burrows at neap tides than during the rainy season (80% and 91%, respectively). However, time spent for feeding activities was similar during both seasons. During almost all observation periods crabs collected leaf litter, but rarely fed on it outside burrows. At neap tides nearly all available litter was collected, suggesting that the U. cordatus population is litter-limited during these times. At spring tides (regular tidal inundation) the surface activity of U. cordatus was tide-dependent. Crabs closed their burrow entrances 2-3 h before flooding and re-emerged as soon as the tide retreated. During the day, burrow maintenance was the second most frequent behaviour after feeding. Agonistic interactions were regularly observed and were mainly related to burrow defence. The mean foraging radius of the crabs was only 19 cm (max: 1 m) underneath high Rhizophora mangle trees where crab densities were high. The results point to a high competition for burrows and show that U. cordatus is territorial. It is concluded that several exogenous factors, in particular light, leaf litter availability, flooding of burrows and the presence of conspecifics are important in controlling the crabs' activity patterns.     

Activity and feeding of Dotilla fenestrata (Brachyura: Ocypodidae) in a warm,temperate South African estuary

The activity of and consumption of organic material by the sand-bubbler crab Dotilla fenestrata was studied over neap and spring tides on a sheltered sand bank close to the mouth of the warm, temperate Kowie Estuary, South Africa. Crabs emerged from their burrows only after the tide receded, and it was light. Time to emergence therefore varied from about 30 minutes to three hours after exposure, depending on the time of low water in the early morning vs at midday. General activity of the crab population was longer on a spring tide (about five hours) than on a neap tide (about three hours). Maximum densities of active crabs were 140 m^–2 and 41 m^–2 on spring and neap tides, respectively. After emergence, crabs spent 60% to 80% of their time feeding. In the Kowie Estuary, D. fenestrata produced between 7 and 12 pseudofaecal pellets, average weight 0.0358 g per pellet, per minute. These pellets had a significantly lower organic and chlorophyll a content than the substratum, and it was estimated that crabs removed about 25% of the organic content from the sediment.     

Semilunar courtship rhythm of the fiddler crab<Emphasis Type="Italic"> Uca lactea</Emphasis> in a habitat with great tidal variation

Semilunar courtship rhythm is a widely distributed phenomenon among fiddler crabs in the genus Uca (Decapoda, Ocypodidae). Typically, synchronous courtship has been reported to peak near spring tides. To determine whether a region of large tidal variation shifts reproductive activity, we measured the frequency of specific courtship behaviors including claw-waving and semidome building for U. lactea males on Kanghwa Island, Korea. We found that synchronized courtship for U. lactea peaked near neap tides, whereas near the spring tides, seawater flooded the habitat and males predominantly fed on the mudflat. Although active females, which hold their burrows and usually feed on the mudflat, are abundant near to spring tides, males rarely claw-waved to attract females. This pattern is atypical for the species because other populations of U. lactea on Japan and Taiwan are synchronous around spring tides. We suggest that males invest most of their time in feeding during spring tides because foraging is limited during neap tides. During neap tides, males feed infrequently and thus expend stored energy on courtship signals. We conclude that patterns of reproductive synchrony may be dependent on food availability in periodically changing environments.     

Tidal amplitude and fish abundance in the mouth region of a small estuary

Using an acoustic underwater camera (Dual Frequency IDentification SONar, DIDSON), the abundance and direction of movement of fishes > 80 mm total length (L_T) in the mouth of a small South African estuary during spring and neap tidal cycles were observed. While the sizes of fishes recorded were consistent across both tide cycles, the number of fishes passing the camera was significantly greater during the smaller neap tides. Schooling behaviour was more pronounced for fishes that were travelling into the estuary compared to fishes swimming towards the ocean.     

Role of the tidal cycle in the gonadal development and spawning of the tropical wrasse Halichoeres trimaculatus

Many wrasses on coral reefs exhibit daily spawning that peaks around daytime high tides. In this study, we examined tidal-related ovarian development in the threespot wrasse, Halichoeres trimaculatus, a species common on coral reefs in the Indo-Pacific Ocean. When the fish were collected in the morning at different tidal phases, the gonadosomatic index (GSI) and ovarian histology changed; concomitant with increases in GSI towards high tide, a clutch of the most advanced oocytes developed from vitellogenic to maturation stages. Ovulated eggs and post-ovulated follicles (POF) existed in most ovaries around high tide, but only POF remained around ebb tide, suggesting that spawning occurred during or after high tide. We noticed that tidal-related spawning was considerable in the morning and that most ovaries collected on the afternoon high tide exhibited post-spawning features. This suggests that certain labrid species possess plasticity with regard to their spawning time and utilize potent environmental cues to ensure their reproductive success. When pieces of ovary were incubated with precursor steroids, high conversion of testosterone to 17beta-estradiol occurred during high and ebb tides, while that of 17alpha-hydroxyprogesterone to 17alpha,20beta-dihydroxy-4-pregnen-3-one and 17alpha,20beta,21-trihydroxy-4-pregnen-3-one was observed during low and flood tides. Incubation of pieces of ovary with human chorionic gonadotropin resulted in similar fluctuations in the steroid hormones with tidal phase. Production of these steroid hormones correlated with oocyte development in the ovaries and was probably regulated by gonadotropin. These results demonstrate that the daily cycle is fundamental for oocyte development, and that the tidal cycle is superimposed on this process.     

Tidal and diurnal variations in larval fish abundance in an estuarine inlet in Ariake Bay,Japan: implication for selective tidal stream transport

We have conducted a preliminary study of tidal and diurnal variations in the distribution of dominant larval and juvenile fishes in the Chikugo River inlet (Ariake Bay, Kyushu, Japan) to determine whether selective tidal stream transport (STST) occurs. Larval and juvenile fish were collected from the mesohaline zone of the Chikugo River inlet during spring 2002. Temperature, salinity, depth, and current velocity were measured. Larval and juvenile abundance were compared among four tidal conditions, flooding tide, high tide, ebbing tide, and low tide, and between day and night. A total of 12 families, 15 species, and 5,577 individuals were collected. Temperature did not vary significantly with tidal conditions whereas salinity, depth, and current velocity varied significantly. Salinity also was correlated significantly and positively with depth. The abundance of most of the fishes was correlated positively and significantly with salinity and depth. Lateolabrax japonicus, Trachidermus fasciatus, Acanthogobius hasta, and other gobiid larvae (Gobiidae spp.) were significantly more abundant during high tide; in contrast, Coilia nasus and Neosalanx reganius were most abundant during low tide. The abundance of most of the fishes was higher during high tides at night than during the day, indicating the existence of STST, which may be strategically associated with ascending progress to upstream nursery areas.     

Feeding habit of larval and juvenile ayu,Plecoglossus altiyelis in the surf zone of Tosa Bay,Japan

A total of 7,000 larval and juvenilePlecoglossus altivelis was collected at semimonthly intervals with a small seine in a surf zone of Tei beach facing Tosa Bay during the period of June 1982 to May 1983. They occurred in the surf zone from middle October to middle May. About 500 larvae and juveniles (10.9–59.9 mm TL) were used to examine their feeding habit. The feeding incidences by collection dates fluctuated from 0 to 100%, with 90.6% in total incidence. They fed mainly on copepods (e.g.Paracalanus parvus andOithona spp.) throughout postlarval and juvenile stages, while they first took small benthic animals at 53.0 mm TL. Their food compositions were influenced fundamentally by the planktonic fauna of the surf zone, but larvae under 20 mm TL tended to take relatively larger copepods.     

Temporal and spatial patterns in the abundance of wintering Red-breasted Mergansers Mergus serrator in an estuary

Tidal, temporal and spatial aspects in the abundance of wintering Red-breasted Mergansers in the Ythan estuary, northeast Scotland, were investigated. The number and location of birds were examined in relation to four variables: tidal cycle and tidal range, and month and time of day. A significant increase in the numbers of this piscivorous bird occurred as the winter progressed. Numbers were highest in early morning and then decreased significantly during the day. Spring tides were favoured over neap tides and a significant increase in numbers was recorded between low tide and high tide. The lower part of the estuary was preferred over the upper part with no detectable shift in the preference occurring in relation to any of the four variables.     

Feeding rhythms of the green-lipped mussel, Perna viridis (Linnaeus, 1758) (Bivalvia: Mytilidae) during spring and neap tidal cycles

Food availability and feeding responses of the green mussel Perna viridis were investigated for two complete tidal cycles during both spring and neap tides. Temporal changes in total particulate matter (TPM), particulate inorganic matter (PIM) and particulate organic matter (POM), were smaller during neap than spring tides. Seston characteristics at different times of a tidal cycle were compared for both spring and neap tides. Only during spring tides were TPM and PIM significantly higher at high tides while POM remained relatively constant (P>0.05). The clearance rate of the mussels underwent temporal variations with tides, and was a negative power function of TPM and a positive linear function of f (organic content), during both spring and neap tides. f was the key factor influencing filtration rate, organic ingestion rate, absorption rate and absorption efficiency. All feeding rates increased linearly with increases in organic content. Pseudofaeces were produced only during spring but not neap tides. Feeding rates and absorption efficiency were highest at low and lowest at high tides (P<0.01). There was no significant temporal change in the wet weight and protein content of the crystalline style with the tidal regime. For the digestive gland, alpha-amylase activity was higher at spring than at neap tides, and higher during high tides in a tidal cycle. The digestive gland cellulase activity did not change significantly with the tides. For the crystalline style, both the activity of cellulase and alpha-amylase were not significantly different (P>0.05) between spring and neap tides. Tidal rhythms in feeding and digestion in this species were likely controlled by temporal variations in food availability in the seawater. By adjusting feeding rates and enzymatic activities, absorption in Perna viridis remained constant, irrespective of the changes in food availability.     

Seasonal occurrence and food habits of larvae and juveniles of two temperate basses in the Shimanto estuary,Japan

Larvae and juveniles ofLateolabrax japonicus andL. latus occurred from January to May 1986 in the shallow waters of the Shimanto estuary.L. japonicus markedly outnumberedL. latus. Distinct ecological differences were recognized in habitats and food habits between the two species:L. japonicus mainly inhabited eelgrass beds composed ofZostera nana, whileL. latus appeared evenly in both eelgrass beds and non-eelgrass habitats; the former fed on copepods and cladocerans, while the latter fed on copepods and fish larvae. From these habitat and food habit analyses, estuaries were considered to be important as a main habitat forL. japonicus, but not forL. latus. The fact that ecological differences have occurred during the early life stages was inferred to be one of the possible keys to speculate on the speciation of the two species.     

Intertidal territoriality and time-budget of the surgeonfish, Acanthurus lineatus, in American Samoa

Synopsis The herbivorous surgeonfish Acanthurus lineatus aggressively maintained feeding territories in the surf zone of the outer reef flat in American Samoa. Intertidal territories were re-established each morning, as well as after displacement by low tides or rough surf. Day-to-day site fidelity of recognizable individuals was high: 99.9% return rate per day for adults (15–20 cm), 99.6% for juveniles (8–13 cm), and 97.2% for recruits (2.5–5 cm). Fish fed on turf algae primarily in the afternoon (80% of available time), and spent 10% of their time on active territorial defense and 2–13% of their time on forays from their territory. On average, a fish defended its territory 1900 times daily and took 17 000 bites (= 7400 bites m^–2 d^–1), but rough surf reduced feeding by 60% and defense by 75%. High territorial defense requirements significantly reduced feeding rates. Although the distribution and behavior of this species in Samoa was in large part similar to that reported for it elsewhere (Australia, Indian Ocean), there were notable differences: in Samoa A. lineatus densities within colonies were greater (0.4 fish m^–2), territory size was smaller (2.3 m²), and defense rate against intruders was greater (2.5 attacks min^–1). These differences in Samoa may be related to their smaller body size, greater abundance or increased food supply caused by hurricane damage to reefs which has enhanced the algal turfs that A. lineatus feeds upon.     

Rapid changes in fish utilization of mangrove habitat in Western Madagascar

Fish use of a mangrove habitat was studied in a small mangrove forest on the West coast of Madagascar. A sand bar near the inlet retains water in parts of the channel (pools) at low tide. Fishes in four of these pools were examined daily at all phases of the tidal cycle for 3 weeks using underwater visual census. During week 1, fishes were diverse and abundant in all pools: the dominant species were cardinalfish (related to Apogon lateralis); monos, Monodactylus argenteus; black spotted snappers, Lutjanus ehrenbergi; double bar bream, Acanthopagrus bifasciatus; emperors, Lethrinus lentjan and L. sp., surgeon fish, Acanthurus nigricauda; red-lined sweetlips, Plectorhinchus plagiodesmis; and butterflyfish, Chaetodon kleini. Some species were more abundant in shaded pools; others in more open pools. During week 2 a dramatic difference was noted: the only fishes found were schools of cardinalfish and one moray eel. This week had neap tides, with high tides in the morning and low tides in the afternoon. As the week progressed and during week 3 (spring tides), fishes slowly repopulated the habitat and diversity increased. Monos, absent in week 2, now had increasing numbers of small individuals. While large emperors were scarce, small individuals appeared. The larger butterflyfish and surgeonfish seen in week 1 were replaced by small ones during week 3. Species that had been rare in week 1 were more abundant, including pipefish and small barracudas. While species richness increased during week 3, the community was strikingly different from that seen 2 weeks earlier. Only Pool 1, closest to the entrance, recovered its original species richness. Abundance was much lower than in week 1. Our snapshot study apparently captured a time when older juveniles left the mangrove forest and smaller fishes recruited into it. Utilization of this habitat will likely vary throughout the year depending on the reproductive cycle of the different species whose juveniles utilize it. Longer studies are needed to learn about cycles in fish use of the mangroves.     

Behavioural basis of intertidal zonation in Eurydice pulchra Leach

Eurydice pulchra Leach freshly collected at neap and spring tides exhibits an endogenous circatidal rhythm of swimming when kept without sand in constant conditions in laboratory aquaria. When provided with sand in the laboratory, tidally-phased swimming may occur spontaneously at some times of the lunar month, predominantly after the times of highest spring tides but not at neaps. In these cases, therefore, there appears to be clearly demonstrated for the first time in a marine animal a circasemilunar pattern of emergence from the substratum which is coupled to a circatidal rhythm of swimming; the isopods in the laboratory emerge from the sediment only when the swimming rhythm and emergence rhythm are at their maxima. In the field at neap tides some isopods emerge to swim at high tide, dependent upon their responses to water agitation, their feeding state and responses to light. At and just after spring tides in the field large numbers of the isopods swim at high tide, triggered by the endogenous circasemilunar rhythm of emergence which in the field is probably cued by the effects of increased water agitation. The main function of the endogenous circatidal rhythm of swimming appears to be to permit the isopods to return to their preferred zone in the sand before the fall of the tide. The coupled circasemilunar rhythm of emergence which induces greater swimming at spring tides reduces the risk of animals being stranded high on the shore during neap tides.     

Evidence for tidally-induced vertical migration of some gelatinous zooplankton in the Wadden Sea area near Sylt

Gelatinous zooplankton is more abundant in the Wadden Sea area of North Sylt than in the adjacent North Sea. The hypothesis is tested that medusae maintain their position in the North Sylt Wadden Sea by ascending to the surface at flood tides and descending to the bottom during ebb tides, thus avoiding the strong ebb currents which could carry them out of this food-rich area. Surface sampling at a main tidal channel revealed great differences between high tide and low tide abundances of five species of medusae in the surface layer.Rathkea octopunctata, Sarsia tubulosa, Eucheilota macultata andPleurobrachia pileus were all more abundant around high tide than during low tide. Bongo net sampling in different depth layers revealed thatPleurobrachia pileus, Bougainvillia ramosa andEucheilota maculata showed a preference for the surface layers around high tide, whereas most of the individuals were found in the deepest layer around low tide. The results suggest tidally-induced vertical migration of medusae in tidal channels. This may assist maintenance of the populations in the Wadden Sea area near Sylt.     

Tidal migration and patterns in feeding of the four-eyed fish Anableps anableps L. in a north Brazilian mangrove

The tidal migration, temporal and spatial patterns in feeding of the surface-swimming four-eyed fish Anableps anableps (Anablepidae) were studied in a macrotidal mangrove area in north Brazil to exemplify the ecology of a tropical intertidal fish. Visual censuses in the main channel showed that abundances were high at low water (LW) and low at high water (HW). Anableps anableps entered the intertidal creeks with the first flood rise. They fed in the inundated mangrove at HW and returned gradually after the ebb current maximum to concentrate again in the subtidal parts of the main channel at LW. This pattern occurred at neap, mid and spring tides throughout the year. The tidal migration was triggered by water level, not by time. In the study area the diet of A. anableps caught with block nets was dominated by intertidal red algae ( Catanella sp.). Other important food items were Insecta and Grapsidae. The combination of high inundation and daylight (spring tide-day) provided the best foraging conditions, probably emphasizing the importance of the above-water eye. Darkness and low inundation was linked to poorest foraging conditions (neap tide-night). The quantity of food consumed by A. anableps was clearly influenced by the factors tide, time of day and creek location, but not by size and sex. The qualitative composition of the diet was not influenced by any of the factors, except for mud, which was only ingested at neap tides. The temporal and spatial variability in food consumption suggests that food web modelling in macrotidal areas may lead to misinterpretations of the overall systems dynamics if the spring and neap tide alternation and the diurnal cycle are not considered.