A quantitative description of the behaviour of wild juvenile plaice (Pleuronectes platessa L.)

The behaviour of young plaice (Pleuronectes platessa L.) as they migrate up and down sandy beaches with the tide is described. Their behaviour during this migration consists mainly of swimming and feeding interspersed with rarer behavioural acts. Over short periods swimming behaviour can be described by a random model in which the probability of a swim occurring remains constant. This probability varies markedly, however, from hour to hour. Two types of swimming movement are recognized: one of very short duration represents searching for food and the other, longer, category serves to transport the fish up and down the shore. Variations in the feeding rate and in the frequency, duration, and direction of swimming movements over the tidal cycle are described and related to the changing physical and biological conditions that the fish experience during their intertidal movements.     

An optimal foraging and migration model for juvenile plaice

Summary A state-dependent model has been used to predict daily and tidal patterns of migration, feeding and inactivity in juvenile plaice (Pleuronectes platessa L.) in their intertidal and shallow subtidal nursery areas. Vertical position, quantity of energy reserves and fullness of the gut characterized the state of individual fishes. If feeding is visually mediated, the model predicts that, by night, plaice should move to areas of low predation risk and become inactive, whereas by day, plaice should migrate to feed in areas of high prey encounter rate. Vertical zones adopted by day and night and, hence, patterns of migration, should depend on the distributions of predators and prey. When prey are more abundant in the intertidal zone, plaice should move onshore to feed as the tide rises and when prey are more abundant offshore, plaice should move offshore to feed by day. If predators are equally abundant in all zones, the fish should behave as if no predators were present, having no effective refuge. An increase in the abundance of predators with depth results in the restriction of plaice activities to shallower vertical zones, depending on the magnitude of the predation threat. Zones adopted thus depend on the trade-off between energy intake and predation risk. Concordance between predicted behaviour and observed patterns is evident in contrasting habitats. Migration and feeding in the Wadden Sea, where prey are more abundant on intertidal flats, is dominated by the tidal component, whereas on impoverished exposed beaches of the west coast of Scotland, the diurnal component is dominant. Tidally related behaviour persists in the latter environment, not predicted by the model and may be a consequence of using endogenous rhythms to approach optimal behavioural patterns.     

Depth selection behaviour during activity cycles of juvenile plaice on a simulated beach slope

Juvenile plaice Pleuronectes platessa on a sand slope in a laboratory tank showed depthselection behaviour consistent with offshore migration by day and onshore by night, as seen in natural conditions. Plaice stayed deep (0·5 m) and avoided the shallowest water (<10 cm depth) during light periods, but ventured up the slope and into the shallows in darkness. The freshly caught fish showed circatidal cycles of activity, but showed no change in depth selection between periods of high (time of expected high water) and low activity (expected low water). This suggests that changing direction of movements in tidal migration is controlled by responses to changing environmental conditions. Further analysis showed movements of fish up the slope to be slower, covering less distance and with shorter pauses in between than moves down the slope. Frequency, distance travelled and speed of moves up and down the slope did not change with tidal or light-dark cycles, and so such modulation could be ruled out as a mechanism for migration.     

Being there and getting there: balancing the costs of migration against the benefits of changing habitat for North Sea plaice

To improve survival and reproductive success, many fish species have evolved migratory life‐histories, showing ontogenetic and/or seasonal changes in habitat use. Individuals move between different areas, each of which is 'best' for a particular activity, such as feeding, growing or spawning. The benefits of moving to a different habitat, however, have to be balanced against the costs of migration, so any behaviour that reduces the cost of movement would be expected to expand the migratory range and thereby increase the range of habitats that can be exploited.
Previous tracking experiments in the southern North Sea have shown that plaice selectively exploit the tidal streams to aid their spawning migration. Here we examine whether this behaviour is primarily (1) an energy‐saving strategy, or (2) a transport mechanism by which fish that are unable to navigate over long distances can be carried reliably between feeding and spawning grounds. Because selective tidal stream transport requires that fish remain stationary on the sea‐bed during the 'non‐transporting' tide, energetic calculations predict that this behaviour is beneficial only when the current speed exceeds a critical, size‐dependent, value. We have used detailed information about migratory behaviour from individual fish to calculate the metabolic costs and the likely reproductive benefits of different migratory strategies. Our results show that plaice use selective tidal stream transport only in areas where the tidal streams are suitably fast, indicating that this behaviour is primarily an energy saving strategy.     

Sex‐biased dispersal in a salmonid fish: implications for growth rate and survival

Migration is widespread among marine fishes, yet little is known about variation in the migration of individuals within localities. We tested the hypothesis that variation in the migratory behaviour among plaice ( Pleuronectes platessa ) in the North Sea could be explained by large‐scale differences in the speed and directions of the tidal streams, which the fish use as a transport mechanism. Towards this end, 752 mature female plaice tagged with electronic data storage tags were released at eight locations with contrasting tidal flow properties, between December 1993 and September 1999. The experiment yielded 20 403 days of data from 145 plaice. The position of each fish was determined at intervals throughout the liberty period using the tidal location method. The results show 3 geographically discrete feeding aggregations during the summer, which dispersed over the southern North Sea and Eastern English Channel to spawn during winter. Our results re‐affirmed the major role of the tidal streams in the southern North Sea in structuring plaice dispersion, both by providing transport and guidance, and by delimiting the extent of distribution due to thermal stratification during the summer. These results confirm the prediction that large‐scale variation in migration behaviour can be explained in part by the tidal guidance and transport mechanisms available. They have revealed features of spatial dynamics not previously observed from a century of conventional tagging experiments and illustrate how the study of individual fish can successfully define the migratory characteristics of populations.     

Tidal stream mediation of metapopulation structure in North Sea plaice

Migration is widespread among marine fishes, yet little is known about variation in the migration of individuals within localities. We tested the hypothesis that variation in the migratory behaviour among plaice (Pleuronectes platessa) in the North Sea could be explained by large‐scale differences in the speed and directions of the tidal streams, which the fish use as a transport mechanism. Towards this end, 752 mature female plaice tagged with electronic data storage tags were released at eight locations with contrasting tidal flow properties, between December 1993 and September 1999. The experiment yielded 20 403 days of data from 145 plaice. The position of each fish was determined at intervals throughout the liberty period using the tidal location method. The results show 3 geographically discrete feeding aggregations during the summer, which dispersed over the southern North Sea and Eastern English Channel to spawn during winter. Our results re‐affirmed the major role of the tidal streams in the southern North Sea in structuring plaice dispersion, both by providing transport and guidance, and by delimiting the extent of distribution due to thermal stratification during the summer. These results confirm the prediction that large‐scale variation in migration behaviour can be explained in part by the tidal guidance and transport mechanisms available. They have revealed features of spatial dynamics not previously observed from a century of conventional tagging experiments and illustrate how the study of individual fish can successfully define the migratory characteristics of populations.     

Effects of solar radiation on the population dynamics of juvenile flatfish in the shallows of the Wadden Sea

During the first days after their arrival in the Wadden Sea, newly settled plaice Pteuronectes platessa L., flounder, Platichthys flesus L., and sole, Solea solea L., stay at low tide in the residual waters on the tidal flats. Here they can be exposed to high levels of solar radiation and extreme temperature conditions. As a result of the extreme temperatures, all settlers eventually leave the shallows and start to exhibit tidally-phased feeding migrations from the tidal channels on to the submerged fiats, In contrast to warm summers, for example 1982, no formation of a hyaline zone could be detected in the otoliths of surviving O-group plaice in 1990, 4 weeks after such a mass emigration, which could be directly observed on particular days in both hot and cold summers. Analysis of UV-B radiation data indicated that in 1990 cloud cover and wind conditions during the settling period protected the post-larvae from high temperature and radiation stress which is necessary to induce hyaline zones in the otoliths. Nevertheless, histological examination of the dorsal skin revealed cellular changes characteristic of sunburn damage in part of-the O-group plaice population in 1990.
Mortality of O-group plaice during and after such a mass emigration, which was directly observed in the drainage gulleys in 1982, was estimated at approximately 14–27%. This rate is assumed to be close to the maximum mortality which can be expected from abiotic factors for O-group plaice in the Wadden Sea.     

Tidal movements of shallow water fishes in Kuwait Bay

The intertidal migrations of shallow water fishes in Kuwait Bay were studied during two 24 h periods using an otter trawl. Most species wereconcentrated at a location corresponding to Low Low Water mark. At the other tidal levels some fishes migrated with the tide to High High Water mark through a horizontal distance of up to 2 km. Any effect of day and night on the catch was masked by the effect of tidal condition.
Leiognarhus decorus (de Vis) was typically concentrated at Low Water but a proportion of the population followed the rising tide toward the shore. There was a die1 effect whichmay have been due to fish moving to shallower water at night compared with the day. Length-frequency distribution changed slightly with tidal state so that larger fish were not captured at High Water during the day. Of the fishes captured at high tide, the largest individuals of L. decorus were captured at a depth of 2.0 m deep. Solea elongata Day did not migrate with tidal fluctuations and was captured in large numbers only at Low Water. Arius tenuispinis Day was captured in shoals at or around Low Water.     

Tidal and diel timing of river entry by adult Atlantic salmon returning to the Aberdeenshire Dee, Scotland

The timing of salmon migration from the estuary of the Aberdeenshire Dee into the river in relation to tidal phase and time of day was studied by combined acoustic and radio-tracking of individual fish and by analysing records of untagged fish from a resistivity fish counter 0.8 km upstream from the tidal limit. Up-estuary movements that led to river entry were predominantly nocturnal and tended to occur during the ebb tide. Penetration into the non-tidal reaches of the river also tended to occur at night, but the timing of salmon movements was no longer significantly associated with tidal phase. The tracking data suggested that the reduction in the strength of the association between salmon migration and tidal phase resulted from variability in rates of progress from the estuary to the river. This variability may have been random, or related to changes in migratory behaviour during entry to fresh water. A simulation of upstream progress by groups of salmon illustrated the decay of the relationship between salmon movements and tidal phase and the advance of the average tidal phase of observed salmon movements with increasing distance of the observation point from where movements were initiated. The magnitude of these effects depended on the average rate of upstream progress and variability in the rate of progress, being greatest when upstream progress was slow and variable. These results highlight a limitation of point observations of migration with regard to identifying environmental stimuli for migration and quantifying their effects.     

Foraging movements of Magellanic Penguins Spheniscus magellanicus in the Beagle Channel, Argentina, related to tide and tidal currents

This study investigates the movements of Magellanic Penguins Spheniscus magellanicus breeding on Isla Martillo during the early chick-rearing period. Foraging paths were reconstructed using GPS loggers that registered the penguins′ geographic position, water temperature and depth at regular intervals. The relationship between penguins′ movements and search strategies, tide and tidal currents were assessed. Mean trip duration was on average 14.7 ± 6.9 h (33% overnight), and the maximum distance reached was 24 ± 10 km. All penguins studied foraged to the east of the colony. We identified three phases based on the sinuosity and speed of the trajectory: transit, central and return. Foraging effort was higher during the central phase, followed by the transit phase, and lower in the return phase. Foraging success, measured as the percentage of time at the bottom during each phase, was also highest during the central phase. In all birds studied, the central phase of the foraging trip took place during ebb tide, and birds travelled to the foraging areas with flow tide running in the same direction of displacement. Our study suggests that penguins take advantage of tidal currents to facilitate their movements to and from the main foraging area, thereby reducing the energy expended. Moreover, we suggest that piscivorous diving birds may enhance their catch rate during ebb tide when fish are more concentrated near the channel bed.     

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.     

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.     

Infection of estuarine fish populations by Cryptocotyle lingua (Creplin)

Fish populations in the Medway Estuary become infected with Cryptocotyle lingua (Creplin) shortly after their inshore migrations. The present study examines infection levels between 1973 and 1975 in four fish species; whiting, Merlangius merlangus (L.), pouting, Trisopterus luscus (L.), plaice, Pleuronectes platessa L. and flounder, Platichthys flesus (L.). Frequency of infection was highest in flounder, followed by pouting, plaice and whiting. Unusually, parasite density was lowest in plaice even though gastropod molluscs (the parasite's first intermediate host) were most common in their diet. Seasonal levels of infection are discussed and prove useful indicators to the migratory movements of individual fish populations. In whiting and pouting, incidence of infection increased with length of host while no relationship between infection and age of host was observed in either flatfish. Metacercariae were not distributed randomly over the body surface; in gadoids they were concentrated above the lateral line while in pleuronectids they were most intense in the fins and gills.     

Behavior of Four Species of Fiddler Crabs,Genus Uca,in Southeast Sulawesi,Indonesia

We studied the behaviors of four species of sympatric fiddler crabs on Kaledupa Island, Indonesia. Species differences in activity level, grooming, burrowing and feeding were related to their habitat and food. Uca chlorophthalmus, living in muddy mangrove areas, were inactive and spent most of the time feeding in place. Females fed 50% faster than males and spent more time feeding. U. vocans was the dominant species at the beach in silty sand and was very active. Its feeding rate was about twice that of the former species, females fed more rapidly than males, and many crabs of both sexes fed in droves at the water’s edge during ebb tides. During ebb tides, they spent most of their time feeding, while at flood tide they engaged in a greater variety of activities, including burrow maintenance. They frequently walked while feeding and interacted aggressively. U. tetragonon lived in a pebbly band along one edge of the beach, by a quay. Their feeding rate was comparable in both sexes and slower than that of U. vocans; they fed largely on filamentous algae growing on the quay, which provides better food, and fed faster during flood tide than ebb tide. They spent more time in waving and other sex-related activities, and were seldom aggressive, except during the week of the full moon. Burrowing activities included placing excavated mud balls some distance away from their burrows and re-arranging them. U. dussumieri inhabited the other end of the beach in muddier substrate. They did not have sex differences in feeding rates and their rate of scooping food into their mouths was slow, but feeding claws made multiple pinches of the substrate, thus accumulating more material in each clawful of food.     

Migratory behaviour of post-smolt Atlantic salmon during initial stages of seaward migration

The movements of 24 hatchery-reared Atlantic salmon Salmo salar smolts, with miniature acoustic transmitters (pingers) implanted surgically, were determined after release in the coastal waters of Passamaquoddy Bay (mean tide range 6 m), New Brunswick, Canada, to describe the first stages of seaward migration. Automated pinger detection at fixed sites, and pinger location and tracking by boat were used. Post-smolts left the release area rapidly, and the majority moved to open waters of the bay within several tidal cycles. Initially, post-smolts moved with a seaward orientation on ebb tides and held positions on flood tides. Their movements into open waters were diurnal, and the timing corresponded with the state of the tide during which they moved through a narrow channel. Post-smolts moved preferentially through this passageway with the aid of the tidal stream. Successful movements out through the channel occurred during ebb tides and any movements back in were during flood tides. Ground speed of fish moving through the channel was 4·2 body lengths s⁻¹ and faster than the tidal stream velocities in the channel. The relative velocity of fish swimming through the channel was 2 body lengths s⁻¹. Post-smolt movement was indicative of active, directed swimming with a reliance on ebb-tide transport for migration through a coastal area with strong tidal currents. Some post-smolts moved seaward directly with no apparent period of acclimation for the transfer to the marine environment, whereas others delayed their departure. These differences in behaviour were probably related to asynchrony in smolting when fish were released.     

0-group flatfish growth conditions on a nursery ground (Bay of Canche, Eastern English Channel)

The settlement and growth conditions of three 0-group flatfish species, sole (Solea solea L.), plaice (Pleuronectes platessa L.) and dab (Limanda limanda L.) in the Bay of Canche, French coast of the Eastern English Channel, were investigated during summer 1997. 0-group dab were the more abundant of the three species (75.4%) followed by plaice (21%) and sole (3.6%). Analyses of fish size distribution indicated that settlement of plaice occurred before sole and dab. Growth of sole was faster than that of plaice, which in turn was faster than that of dab (0.68, 0.43 and 0.33 mm d^?1 respectively). Sole differed from plaice and dab by their larger mean length attained at the end of their first summer stay in the nursery ground. Growth performances of 0-group sole and plaice were analysed by comparing estimated growth in the field with predicted maximum growth according to temperature-growth rate models from experimental studies of growth with unlimited food supply. The data of this study suggest that plaice growth may have been limited during late summer, whereas sole growth does appear to have been determined only by water temperature.     

Otolith oxygen and carbon stable isotopes in wild and laboratory-reared plaice (Pleuronectes platessa)

The relationship between water temperature, growth rate, and otolith isotopic ratios was measured for juvenile plaice (Pleuronectes platessa) reared at two temperatures (11 and 17°C) and two feeding regimes (1 and 3 prey items·ml^?1). The otolith isotope ratios in individual fish ranged from ?2 to ?4 for carbon isotope ratios (δ¹³C) and from 0.2 to 1.9 for oxygen isotope ratios (δ¹⁸O). The otolith oxygen isotope ratios were significantly affected by water temperature, but not by feeding level, and there were no significant synergistic effects. The fractionation of oxygen isotopes during otolith growth was independent of individual growth rate. Carbon isotope ratios were not significantly affected by food ration or water temperature, but were related to fish growth rate. The carbon isotope ratios were negatively correlated with fish length in the colder water treatments, and tended to increase with fish length in the warm water treatments. The laboratory-determined relationship between otolith oxygen isotope ratio and water temperature was applied to individuals of five species (plaice, cod, whiting, haddock, gurnard) collected in a single trawl sample. The otolith derived temperatures often overestimated measured water temperatures. The difference between real and estimated water temperatures varied between species, and the closest fit was for field-caught plaice.     

Nocturnal feeding by Dark-bellied Brent Geese Branta bernicla bernicla

The occurrence and extent of nighttime feeding by Dark-bellied Brent Geese Branta bernicla bernicla on salt marshes adjacent to their tidal roost site were assessed on the north Norfolk coast using position-sensitive radio transmitters. Nine birds were monitored on 143 bird-nights, and feeding was recorded on 87. The frequency of feeding by night varied widely between individuals. The mean proportion of the night spent feeding for one bird was 19.7 ± 8%. Of this, 59% occurred within ± 1.5 h of high tide. Neither the proportion of the whole night spent feeding nor the intensity of feeding around high tide varied with the length of the night or with time after sunset at which high tide occurred. The extent of nocturnal feeding was also independent of the brightness and duration of moonlight. The birds fed for significantly longer on colder nights, and there was a significant positive relationship between the proportion of the night spent feeding and maximum temperature the preceding day. Analysis of daytime activity budgets on inland pastures showed that the proportion of time engaged in nonfeeding, energy-expensive activities was also positively related to maximum daytime temperature. Increases in the proportion of the night spent feeding after warmer days were achieved by increasing the period of time spent feeding on either side of the high tide, not by increasing feeding intensity at high tide. It is concluded that the geese compensate for increased energy expenditure during the day by increasing the extent to which they feed at night. Implications of increasing energy expenditure as a result of disturbance during the day are discussed.     

Physiological preparedness and performance of Atlantic salmon Salmo salar smolts in relation to behavioural salinity preferences and thresholds

This study investigated the relationships between behavioural responses of Atlantic salmon Salmo salar smolts to saltwater (SW) exposure and physiological characteristics of smolts in laboratory experiments. It concurrently described the behaviour of acoustically tagged smolts with respect to SW and tidal cycles during estuary migration. Salmo salar smolts increased their use of SW relative to fresh water (FW) from April to June in laboratory experiments. Mean preference for SW never exceeded 50% of time in any group. Preference for SW increased throughout the course of smolt development. Maximum continuous time spent in SW was positively related to gill Na⁺, K⁺‐ATPase (NKA) activity and osmoregulatory performance in full‐strength SW (measured as change in gill NKA activity and plasma osmolality). Smolts decreased depth upon reaching areas of the Penobscot Estuary where SW was present, and all fish became more surface oriented during passage from head of tide to the ocean. Acoustically tagged, migrating smolts with low gill NKA activity moved faster in FW reaches of the estuary than those with higher gill NKA activity. There was no difference in movement rate through SW reaches of the estuary based on gill NKA activity. Migrating fish moved with tidal flow during the passage of the lower estuary based on the observed patterns in both vertical and horizontal movements. The results indicate that smolts select low‐salinity water during estuary migration and use tidal currents to minimize energetic investment in seaward migration. Seasonal changes in osmoregulatory ability highlight the importance of the timing of stocking and estuary arrival.     

Movements of lumpsucker females in a northern Norwegian fjord during the spawning season

The lumpsucker Cyclopterus lumpus is distributed throughout the North Atlantic Ocean and migrates considerable distances between offshore feeding areas and shallow inshore spawning grounds. The number of the lumpsucker has declined since the mid 1980s, probably as a result of overexploitation. The lumpsucker is the preferred host of the sea louse Caligus elongates, which is a problem for marine aquaculture. However, little is known about the biology of the lumpsucker. The aims of the study were to 1) examine the movements of female lumpsucker during the spawning migration, and 2) assess the potential for lumpsucker to act as a vector for transmission of parasites and diseases between aquaculture farms and wild fish. Twenty female lumpsuckers tagged with acoustic transmitters were released during the spawning season in the inner part of Øksfjord, northern Norway and their distribution was recorded by 22 automatic acoustic receivers. The average time until departure from the fjord was 3 days, and within 1 week all fish had left the fjord. Timing of departure from the fjord was unrelated with either tidal current patterns or the time of the day. A high proportion of the fish (75%) were recorded within 200 m of fish farms, but they did not stay for extended periods at these farms. Our results suggest that mature female lumpsucker exhibit a movement pattern characterized by rapid fjord-scale migrations during the spawning season, and that they are not attracted to salmon farms in the same way as a range of other fish species.