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.     

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.     

Contrasting diel activity and feeding patterns of four instars of Rhyacophila dorsalis (Trichoptera)

1. Ontogenetic shifts in prey choice and predator behaviour can affect food‐web structure. Therefore, it is important to establish if the diet and feeding activity differ between life‐stages of the same species. This hypothesis was tested for second, third, fourth and fifth larval instars of Rhyacophila dorsalis by comparing their diel activity and feeding patterns. Second to fifth instars collected from two streams were used either for gut analyses or for observations of their activity and feeding patterns in three stream tanks. Food was provided in excess; being organisms living in bryophytes on top of a large stone in each tank, augmented by different‐sized larvae of Ephemeroptera, Simuliidae and Chironomidae. As few first instars for gut analyses were found in the field, the diet of first instars reared in the laboratory was also studied. 2. Larvae for gut analyses were taken 1 h before dusk or dawn (n = 50 larvae per instar for each day or night sample). First and second instars fed on the smaller food items with no significant day‐night differences in diet. Gut contents indicated a progressive trend from feeding chiefly at night in third instars to almost exclusively at night in fifth instars. Fourth and fifth instars fed on the larger food items, whilst the diet of the third instar larvae overlapped with that of both the earlier and later instars. 3. Diel activity patterns of single larvae differed between instars but not within each instar (n = 20 larvae per instar). Second instars were active throughout the 24 h, with peaks at dusk, around midnight, dawn and around midday. A similar pattern was shown by third instars but the peak of activity at midday was less than the other three peaks. Prey were captured only during these peaks for both instars. Fourth and fifth instars were most active, and fed only, at night. They used an ambush strategy to capture more active prey at dusk and dawn (e.g. Baetis, Gammarus), and a searching strategy to capture more sedentary prey during the night (e.g. chironomids, simuliids). These experiments provided support for the hypothesis under test. If competition and/or interference occur between instars, then it could be reduced between earlier and later instars because of differences in their diet and diel pattern of feeding activity.     

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.     

Endogenous rhythm in activity of an estuarine amphipod,Talorchestia longicornis

The biological rhythm in activity of the supratidal amphipod Talorchestia longicornis Say was determined under constant conditions. Surface activity was monitored with a time-lapse video system under red light and assessed as the number of animals emerging from their burrows and active on a sand substrate at 0.5?h intervals. The amphipods had a circadian rhythm in which they were active at the time of night at the collection site. The rhythm could be entrained by a light?:?dark cycle and had an average free-running period of 23.7?h. Activity also appeared to be related to tidal times at the collection site because, under constant conditions, surface activity was suppressed at the time of nocturnal high tide and increased at the time of nocturnal low tide. The rhythm is functionally significant for foraging at night to avoid visual predators, desiccation and high temperatures.     

Endogenous rhythm in activity of an estuarine amphipod, Talorchestia longicornis

The biological rhythm in activity of the supratidal amphipod Talorchestia longicornis Say was determined under constant conditions. Surface activity was monitored with a time-lapse video system under red light and assessed as the number of animals emerging from their burrows and active on a sand substrate at 0.5 h intervals. The amphipods had a circadian rhythm in which they were active at the time of night at the collection site. The rhythm could be entrained by a light : dark cycle and had an average free-running period of 23.7 h. Activity also appeared to be related to tidal times at the collection site because, under constant conditions, surface activity was suppressed at the time of nocturnal high tide and increased at the time of nocturnal low tide. The rhythm is functionally significant for foraging at night to avoid visual predators, desiccation and high temperatures.     

Strong zonation of benthic communities across a tidal freshwater height gradient

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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.     

The influence of habitat, season and tidal regime in the activity of the intertidal crab Neohelice (=Chasmagnathus) granulata

The activity pattern of intertidal crabs is influenced by factors that usually change rhythmically following tidal and/or diel cycles, and is often associated with the use of refuges. The movement activity of the burrowing crab Neohelice granulata was compared among three populations from SW Atlantic coastal areas where they face different tidal regimes, water salinities, substrata and biological factors. At each site, we examined the seasonal activity of the crabs (individuals collected in pitfall traps) in two types of habitat: mudflat and salt marsh. The working hypothesis is that the activity would vary according to the diverse environmental conditions encountered at geographical and local scales. Crab activity varied between sites and seasons showing to be more intense when habitats were covered by water. The most active groups were large males, followed by large non-ovigerous females. Ovigerous females were almost inactive. Most crabs were near or inside burrows at low tides in Mar Chiquita and Bahía Blanca, but they were active at both low and high tides in San Antonio during spring and summer. N. granulata were active in a wide range of temperatures: from 10 to 37 °C at low tides and at temperatures as low as 2 °C when covered by water. Differences of activity between mudflat and salt marsh varied among sites depending on flooding frequencies. Movement activity of N. granulata varied both in space and in time; crabs move under very different abiotic conditions (e.g., low or high tide, daylight or night, low and high temperature) and their movement may also be prevented or elicited by biotic conditions like burrow complexity, food quality and predation pressure. The wide set of conditions under which N. granulata can be active may explain why this is the only semiterrestrial crab inhabiting latitudes higher than 40°S in South America.     

Diel activity and home range size in relation to food supply in a drift-feeding stream fish

Individuals within a species often compete for resources inboth space and time. In dominance hierarchies individuals withthe greatest competitive ability will occupy prime locationsduring optimal periods. We compared spatial and temporal useof habitat, aggression, and feeding success among giant kokopu(Galaxias argenteus) in dominance hierarchies under normal andreduced food supply. Under normal food supply conditions dominantfish were predominantly nocturnal and maintained large homeranges at night. Conversely, subdominant fish were diurnal andoccupied large home ranges by day but were generally not observedat night. When the food supply was limited, the utilizationof pools was determined by social rank. Dominant fish from eachpool increased diurnal activity, home range size, aggressiveness,and the capture of food items offered by day while simultaneouslyreducing the spatial and temporal activity and habitat use ofsubdominant fish. These results indicate that behavioral changesin large dominant fish influence and reduce the amount of resourcesavailable to subdominant fish.     

Patterns of temporal occurrence of brachyuran crab larvae at Saco mangrove creek, Inhaca Island (South Mozambique): implications for flux and recruitment

Temporal patterns of larval occurrence of brachyuran taxa weredescribed from Saco mangrove creek, Mozambique, based on planktonsamples. Brachyuran larvae were collected hourly in four 24h cycles during neap and spring tidal periods at a fixed station,in November 1997 and February 1998. Results indicate a semilunarcycle of larval release activity for most species. Newly-hatchedstages of sesarmids showed a peak occurrence in post-crepuscularebbing tides, Macrophthalmus spp. and Uca spp. showed highestdensities during ebbing tides at night. Dotilla fenestrata andPinnotheridae, in spite of showing a semilunar pattern, didnot display significant differences between day and night. Mostmangrove taxa showed thus a clear larval exportation behaviourduring ebb tides, with little return during the following floods,with varying degrees of dependence on the diel period. Megalopaewere restricted to the night period, and highest values wereobtained during spring tidal periods. However, maximum valuesof megalopae were obtained both during flood and ebb periods,contrary to most previous studies. This is interpreted as abi-directional transport of non-competent megalopae into andfrom the mangrove area.     

The role of site, habitat, seasonality and competition in determining the nightly activity patterns of psammophilic gerbils in a centrifugally organized community

Major ecological forces affecting diel activity patterns include predation, competition, resource dynamics, and ambient conditions. In this work we studied if, how, and why the nocturnal activity patterns of two gerbil species differed with respect to site, habitat, season, and inter-specific competition. The study system included two gerbil species- Gerbillus pyramidum and G. andersoni allenbyi in a sandy landscape composed of three habitats-shifting, semi-stabilized, and stabilized sand habitats. Conforming to previous studies, in the semi-stabilized sand habitat clear temporal partitioning in activity times occurred, with G. pyramidum being active earlier in the night and G. a. allenbyi being active in the later part of the night. As shown in previous studies in a different, more xeric site, removal of G. pyramidum demonstrated that this activity pattern is the result of interference competition by the latter on the former. Nocturnal activity pattern differed between habitats and seasons. Temporal partitioning occurred in the semi-stabilized sand habitat where both gerbil species were common. In the shifting sand habitat, G. pyramidum activity pattern was similar to that in the semi-stabilized sand habitat, but at a lower level. In the stabilized sand habitat where G. a. allenbyi occurred alone, its activity was the highest among the three habitats and was fairly constant throughout the night. Temporal partitioning occurred in the fall and early summer, but not in the early spring. The roles of predation risk, competition and resource dynamics in determining the gerbils' temporal activity together with their implications for community structure, are discussed.     

Short-term changes in the mesozooplanktonic community in theSeine ROFI (Region of Freshwater Influence)(eastern English Channel)

Discrete-depth, hourly mesozooplankton samples were collectedover a 92 h period in May 1992 at an anchor station within theSeine Region of Freshwater Influence (ROFI) (English Channel).The mesozooplankton community defined as a euryhaline marineassemblage was dominated by the calanoid copepods Acartia spp.,Temora longicornis and Centropages hamatus, the cladoceran Evadnenordmanni and the appendicularian Oikopleura dioica. The semi-diurnaltidal current was the dominant factor in determining the short-termtemporal changes in the community in terms of density and speciescomposition so that zooplankton patches displayed oscillatingmotion in relation to tidal advection. Although a few species(e.g. Pleurobrachia pileus) exhibited higher densities aroundlow tide, maximum densities were observed for most species (e.g.T.longicornis and E.nordmanni) around high tide, according tosalinity variations. Diurnal changes were only reported forcyclopoid copepods (i.e. Halicyclops sp. and Cyclopina sp.)which wer$$$ mainly endobenthic during the day and moved intothe water column at night. Besides temporal changes in depth-averageddensities, most species exhibited vertical migrations at dieland/or tidal periods. Tidal vertical migrations were reportedonly for a few taxa and could be the result of passive mechanisms(e.g. vertical mixing) rather than of active behaviour. Dielvertical migrations were observed in most of the abundant taxa.While this migration pattern did not appear to be an adaptationto predator avoidance within the Seine ROFI, it could regulatehorizontal transport of organisms and promote their retention.The consequences of the short-term mesozooplankton fluctuationsfor sampling designs are discussed.     

Studies on activity pattern of the patellid limpet Cellana toreuma (Reeve)

The habits and activity of Cellana toreuma (Reeve) have been studied in a field population. Activity is very high at ebb and flood tide, independent of the time of day, and sharply decreases during complete submergence or complete exposure. Duration of movement fluctuates with tidal amplitude; when neap tide, movement does not stop during low tide owing to continuous awash conditions. Vertical movement entirely coincides with tidal direction at day and night. In this way, the movement does not depend on light but chiefly the tide. Compared with a nocturnal limpet, C. nigrolineata (Reeve), the divergence in the behavioural patterns may be a function of the extent to which limpets are influenced by light. Duration of movement and distance increase with limpet size. Homing behaviour was not observed.     

Diel activity patterns of Carabidae,Staphylinidae and Perimylopidae (Coleoptera) at South Georgia,Sub-Antarctic

Summary Six of the eight species of beetles at South Georgia, the southernmost beetles of the world, were investigated in outdoor arenas with pitfall traps for their diel patterns of locomotory activity. All of them were clearly nocturnal, the only exception being a small staphylinid which appeared to be active throughout the day and night. Activity of the other species was restricted to the dark period of the night, with little activity during twilight. Peak activity occurred before or at midnight. Activity was clearly correlated with temperature. Diurnal activity may involve a risk of overheating and desiccation, but could also have evolved in the past due to bird predation, which today is very low in the study area.     

Habitat use and movements of plains zebra (Equus burchelli) in response to predation danger from lions

Prey species must adapt their behavior to avoid predation. Asa key prey item for lions (Panthera leo), plains zebras (Equusburchelli) were expected to respond to immediate threats posedby lions in their area. In addition, zebras were predicted toexhibit behavior tuned to reduce the potential for encounterswith lions, by modifying their movement patterns in the timesof day and habitats of greatest lion danger. We studied a populationof approximately 600 plains zebra living in Ol Pejeta Conservancy,Kenya. We found that zebra abundance on or near a grasslandpatch was lower if lions had also been observed on that patchduring the same day. Predation danger was highest in grasslandhabitat during the night, when lions were more active. Zebrasightings and global positioning system radio collar data indicatedthat zebras also reduced their use of grassland at night, insteadusing more woodland habitat. Zebras moved faster and took sharperturns in grassland at night. It is hypothesized that these moreerratic movements assist zebras in avoiding detection or captureby lions.     

Temporal Links in Daily Activity Patterns between Coral Reef Predators and Their Prey

Few studies have documented the activity patterns of both predators and their common prey over 24 h diel cycles. This study documents the temporal periodicity of two common resident predators of juvenile reef fishes, Cephalopholis cyanostigma (rockcod) and Pseudochromis fuscus (dottyback) and compares these to the activity and foraging pattern of a common prey species, juvenile Pomacentrus moluccensis (lemon damselfish). Detailed observations of activity in the field and using 24 h infrared video in the laboratory revealed that the two predators had very different activity patterns. C. cyanostigma was active over the whole 24 h period, with a peak in feeding strikes at dusk and increased activity at both dawn and dusk, while P. fuscus was not active at night and had its highest strike rates at midday. The activity and foraging pattern of P. moluccensis directly opposes that of C. cyanostigma with individuals reducing strike rate and intraspecific aggression at both dawn and dusk, and reducing distance from shelter and boldness at dusk only. Juveniles examined were just outside the size-selection window of P. fuscus. We suggest that the relatively predictable diel behaviour of coral reef predators results from physiological factors such as visual sensory abilities, circadian rhythmicity, variation in hunting profitability, and predation risk at different times of the day. Our study suggests that the diel periodicity of P. moluccensis behaviour may represent a response to increased predation risk at times when both the ability to efficiently capture food and visually detect predators is reduced.     

Temporal changes in food preferences of wood mice (Apodemus sylvaticus L.)

Diet choice was determined for wild-caught wood mice (Apodemus sylvaticus L.), temporarily confined to cages in the field and offered a choice of 24–26 types of seeds and fruits in 2-h sessions throughout the night. The mice showed an overall preference for some foods over others. The set-up minimized influences on food preferences of predation risk, hunger, food availability and competition. Variation in food preferences was not attributable to differences between individuals, but followed a temporal pattern. The variety of foods eaten showed a bimodal pattern with peaks corresponding to the two most active periods at the beginning and end of the night. Both the amount of food eaten and variation in the amount diminished from the first to the second active period. An expected selection for carbohydrates early in the night and proteins at the end of the night was not found, but sugars were selected for early in the night. These results are discussed in relation to the conflict between an animal's continuous energy requirements and the essentially periodic activity of foraging.     

Short communication. Food availability and diel feeding rhythms in the marine copepods Acartia grani and Centropages typicus

The effects of food availability on the daily feeding pattern of planktonic copepods was studied for two species: Acartia grani (cultured and wild specimens) and Centropages typicus (wild specimens). Both species showed clear diel feeding rhythms at high food concentrations, with significantly higher clearance rates during night hours. However, at limiting food conditions, while C.typicus maintained its daily feeding pattern, A.grani fed continuously, with similar day and night clearance rates. These results could be explained in relation to differences in predation risks and probability of experiencing starvation episodes in their respective habitats, as well as their capacity for dampening food fluctuations. On the other hand, the maintenance (although with less intensity) of the diel feeding rhythm in A.grani, when cultured in the laboratory for long periods of time (>10 generations) and in the absence of predators, suggests the importance of the endogenous control of copepod feeding patterns.      

Overriding the oddity effect in mixed-species aggregations: group choice by armored and nonarmored prey

Because "odd" individuals often suffer disproportionately highrates of predation, solitary individuals should join groupswhose members are most similar to themselves in appearance.We examined group-choice decisions by individuals in armoredand nonarmored species and predicted that either (1) the oddityeffect would result in preference for conspecific groups forsolitary individuals of both species, or (2) individuals inthe armored species would prefer to associate with groups containingindividuals of the more vulnerable species. Armored brook sticklebacks(Culaea inconstans) and nonarmored fathead minnows (Pimephalespromelas) have the same predators and often occur together instreams. In mixed-species shoals, yellow perch (Perca flavescens)attacked minnows earlier and more often than sticklebacks. Wetested whether solitary minnows and sticklebacks preferred toassociate with conspecific or heterospecific shoals under conditionsof both low and high predation risk. When predation risk washigh, minnows preferred to associate with conspecifics overheterospecifics, as predicted by the oddity effect. In contrast,sticklebacks preferentially associated with groups of minnowsover groups of conspecifics when predation risk was high. Whenpredation risk was low, solitary individuals of both speciespreferentially associated with conspecific over heterospecificshoals. Stickleback shoal choices under low-risk conditionsmay have been influenced by interspecific competition for food.In feeding experiments, minnows were more efficient foragersthan sticklebacks, so it should benefit sticklebacks to avoidminnows unless predation risk is high. Therefore, for armoredprey, the benefits of associating with more vulnerable preyappear to override the costs of both the oddity effect and foodcompetition when predation risk is high.