Space use and feeding patterns of an offshore fish assemblage in a shallow mesotrophic lake

Synopsis Diel and spring/summer space-use and feeding patterns were investigated in an assemblage dominated by five fish species occupying the offshore waters of Lake Opinicon, a shallow mesotrophic lake in southeastern Ontario. We assessed fish distribution and diel movement in May and July through the use of gill nets set at various depths in 1.5–7.0 m depth contour zones, supplemented by observations of fish reaction to the nets. Golden shiners and alewives occupied the upper part of the water column, with the former concentrated at the littoral zone-open water interface, and the latter in the open water. Yellow perch occupied the lower part of the water column in all depth contours. Bluegills were abundant in the upper to midwater depths in all contour zones; black crappies were concentrated in the 2.5–3.5 m zones. All of these species showed either a diel or a spring-summer change in distribution pattern. Bluegills were more abundant in offshore locations in July, whereas golden shiners and yellow perch were more abundant onshore in May. Alewives and black crappies showed distinct diel movements in July, as they were largely absent from the study area during the day, but returned at night to feed. In general, there was more spatial separation among the five species in July than in May.Patterns of spatial distribution among the species generally corresponded with the type and variety of prey consumed, and with diel movement of prey in the case of water column feeders. Other factors that apparently affected spatial distribution and seasonal shifts in this assemblage were risk of predation (golden shiner), spawning activity (alewife), and a decline in prey abundance from spring to summer (bluegill and yellow perch).     

Why are intertidal snails rare in the subtidal? Predation, growth and the vertical distribution of Littorina littorea (L.) in the Gulf of Maine

The North Atlantic gastropod Littorina littorea exhibits a characteristic “intertidal” distribution: the snail is abundant in the littoral zone but scarce in the shallow subtidal and the relatively few subtidal individuals are larger (in shell size) on average than those in the intertidal zone. For highly mobile species like L. littorea, this vertical distribution is primarily determined by directional movement. Biotic and abiotic factors vary across tidal heights, and natural selection for movement to shore levels where fitness is maximized provides the ultimate (evolutionary) explanation for vertical distribution patterns. In this study, we asked whether variation in growth rate and/or predation pressure among tidal heights provide an ultimate explanation for vertical gradients in L. littorea size and abundance. We used a cage experiment to compare juvenile growth rate among tidal heights and a series of field and laboratory experiments to examine variation in predation pressure among tidal heights and snail size classes. Juvenile growth rates were highest in the low intertidal zone, declining at both higher and lower levels. Predation risk for tethered L. littorea increased with both decreasing tidal height and decreasing body size (shell height). Almost all tethered prey were consumed by shell- breaking predators and a census revealed that the two most abundant such predators were the crabs Carcinus maenas and Cancer borealis. Laboratory feeding experiments were used to compare size-dependent prey vulnerability and prey-size preferences for these two key predators. We found that L. littorea vulnerability decreased with increasing snail size and increased with increasing size of both predator species. However, whereas C. borealis were capable of consuming even the largest L. littorea, most Carcinus were unable to feed on individuals larger than 10 mm in shell height. Additionally, C. borealis preferred larger sizes of L. littorea than did Carcinus. Thus, Carcinus, which co-occurs with L. littorea in the intertidal, is a much less effective predator than C. borealis, which is found primarily in the subtidal. We conclude that predation on L. littorea by C. borealis and other subtidal consumers has resulted in the scarcity of this ecologically important grazer in the subtidal. This effect has been produced both through direct predation and by imposing strong selection for movement of L. littorea to higher tidal zones.     

The intertidal movements and distribution of young fish on a sandy beach with special reference to the plaice (Pleuronectes platessa L.)

The intertidal movements of young fish were studied on a sandy beach over several tidal cycles during the day and the night. Young plaice (Pleuronectes platessa L.), by virtue of their abundance, received the most attention, and it was found that they migrate up and down the beach with each tide in company with several other less abundant species. The dab (Limanda limanda (L.)), does not make this migration.The vertical distribution of the commoner fish species was charted and each was found to occupy a specific zone in relation to the water, but with large overlaps in many cases. A close relationship between the length of plaice and dabs and the depth in which they were caught was evident during the summer months, but this relationship broke down in the winter due mainly to an offshore movement of larger fish.The food intake of plaice was studied throughout the day and night; feeding occurred mostly during the day. No effect of the tide on feeding could be detected.The possible mechanisms controlling the tide-related movements and the size-depth relationship are discussed and a general hypothesis is put forward in which it is proposed that the fish are able to select and maintain their position at a particular depth throughout a tidal cycle thereby being guided up and down the beach. Attributes of depth which the fish might use as a reference point in their movements are discussed.     

Sequential movement into coastal habitats and high spatial overlap of predator and prey suggest high predation pressure in protected areas

In theory, predators should attempt to match the distribution of their prey, and prey to avoid areas of high predation risk. However, there is a scarcity of empirical knowledge on predator and prey spatial use when both are moving freely in their natural environment. In the current study, we use information collated on a predators’ diet, its population structure, as well as predator and prey relative abundance, and track the movements of predator and prey simultaneously to compare habitat use and evaluate predation pressure. The study was conducted in elasmobranch protected areas of coastal Tasmania, Australia. The species considered were the broadnose sevengill shark Notorynchus cepedianus, the apex predator in the area, and five chondrichthyan prey species. Notorynchus cepedianus and its prey show similar seasonality in the use of these coastal areas: more abundant in warmer months and absent in winter. Predator and prey also showed high spatial overlap and similar habitat use patterns. These similar movement patterns of predator and prey combined with the additional ecological information (diet, population structure of predator, relative abundance of predator and prey) suggests that N. cepedianus move into coastal areas to exploit seasonally abundant prey. Also, while in protected areas, chondrichthyans are subjected to high predation pressure. Overall, results illustrate the value of simultaneously recording and integrating multiple types of information to explore predator–prey relationships and predation pressure.     

Effects of chlorogenic acid-and tomatine-fed caterpillars on the behavior of an insect predator

If generalist insect predators are a selective force contributing to patterns of feeding specialization by insect herbivores, then predators should be deterred from eating allelochemical-fed prey. The attack and feeding behaviors of naive predators (Podisus maculiventris stinkbugs) reared on control caterpillars (Manduca sexta) fed plain diet were compared to experienced predators reared on caterpillars fed tomato allelochemicals. Tomatine-fed prey were found more quickly by both naive and tomatine-experienced predators, and chlorogenic acid-experienced predators were more stimulated to begin searching for prey. However, experienced predators were less likely to attack both chlorogenic acidfed and tomatine-fed caterpillars than were naive predators. These results indicate that allelochemical-fed prey were easier for predators to locate, but allelochemical-containing prey often deterred predation by experienced predtors.     

Fish community structure and food chain dynamics in the surf-zone of sandy beaches: The role of detached macrophyte detritus

The surf-zones of sandy beaches near Perth, Western Australia often harbour huge accumulations of detached macrophyte detritus. During 2.5 yr sampling, 29 species of fishes were captured over two sandy beaches in this region and the fish community was dominated by juveniles. There was a highly significant positive relationship between the number of fishes and the quantity of detached macrophytes taken in each surf-zone netting. Comparisons of total fish abundance on beaches with and without surf-zone accumulations of detached plants, showed that fishes were two to 10 times more abundant on the beach with weed accumulations, depending on the time of day, and date of sampling. However, despite the overall lower abundance of fishes on the open sandy beach, there was a significant increase in the number of fishes captured over the sandy beach at night. There were also two to five times the number of species over the beach with weed during the day, as opposed to equal numbers of species at night. Seven fish species made up >95% of the total catch and these species fell into two groups with regard to diurnal distribution patterns; those that were equally abundant in weed dominated or open surf-zones, and those that were weed-associated. Analyses of the diets of these fishes and the daytime distribution of an important avian piscivore in the surf-zone suggested that the large quantities of weed in the surf-zone of sandy beaches in this region provide both a rich feeding site for fishes, as well as a refuge from diurnal predators. At night, when visual feeding predators are absent, some fish species move to open sandy areas to feed. Because the majority of fishes in this surf-zone community feed on weed-associated prey, and the input of macrophyte detritus is the major source of primary production in the surf-zone, we argue that the food chain dynamics in the surf-zone in this region are fundamentally different to those of sandy beaches that have been studied previously.     

Predation by gulls on crabs in rocky intertidal and shallow subtidal zones of the Gulf of Maine

Most organisms in intertidal areas are marine in origin; many have distributions that extend into the subtidal zone. Terrestrial predators such as mammals and birds may exploit these animals during low tide and can have considerable effects on intertidal food webs. Several studies have shown that avian predators are capable of reducing densities of sessile and slow-moving intertidal invertebrates but very few studies have considered avian predation on mobile invertebrate predators such as crabs. In this study, we investigated predation by Great Black-backed Gulls (Larus marinus Linnaeus) on three species of crabs (Cancer borealis Stimpson, Cancer irroratus Say, and Carcinus maenas Linnaeus). The study was at Appledore Island, ME (a gull breeding island) and 8 other sites throughout the Gulf of Maine, including breeding islands and mainland sites. On Appledore Island, intertidal and subtidal zones provided over one-third of prey remains found at gull nests, and crabs were a substantial proportion (∼ 30% to 40%) of the total remains. Similarly, collections of prey remains from intertidal areas indicated that crabs were by far the most common marine prey. C. borealis was eaten far more often and C. irroratus and C. maenas less often than expected at each site. Comparing numbers of carapaces to densities of crabs in low intertidal and shallow subtidal zones at each site, we estimated that gulls remove between 15% and 64% of C. borealis during diurnal low tides. The proportion of C. borealis eaten by gulls was independent of proximity to a gull colony. Approximately 97% of the outer coast of Maine is within 20 km of a breeding island. Thus, a lot of gull predation on crabs may occur throughout the Gulf of Maine during summer months. Crabs are important predators of other invertebrates; if predation by gulls reduces the number of crabs in intertidal and shallow subtidal areas, gulls may have important indirect effects on intertidal food webs.     

Wait Until Dark? Daily Activity Patterns and Nest Predation by Snakes

Predation involves costs and benefits, so predators should employ tactics that reduce their risk of injury or death and that increase their success at capturing prey. One potential way that predators could decrease risk and increase benefits is by attacking prey at night when risks may be reduced and prey more vulnerable. Because some snakes are facultatively nocturnal and prey on bird nests during the day and night, they are ideal for assessing the costs and benefits of diurnal vs. nocturnal predation. We used automated radiotelemetry and cameras to investigate predation on nesting birds by two species of snakes, one diurnal and the other facultatively nocturnal. We predicted that snakes preying on nests at night should experience less parental nest defence and capture more adults and nestlings. Rat snakes (Pantherophis obsoletus) were relatively inactive at night (23–36% activity) but nearly always preyed on nests after dark (80% of nest predations). Conversely, racers (Coluber constrictor) were exclusively diurnal and preyed on nests during the times of day they were most active. These results are consistent with rat snakes strategically using their capacity for facultative nocturnal activity to prey on nests at night. The likely benefit is reduced nest defence because birds defended their nests less vigourously at night. Consistent with nocturnal predation being safer, rat snake predation events lasted three times longer at night than during the day (26 vs. 8 min). Nocturnal nest predation did not make nests more profitable by increasing the likelihood of capturing adults or removing premature fledging of nestlings. The disconnect between rat snake activity and timing of nest predation seems most consistent with rat snakes locating prey during the day using visual cues but waiting until dark to prey on nests when predation is safer, although designing a direct test of this hypothesis will be challenging.     

The effect of sand and light on predation of juvenile plaice (Pleuronectes platessa) by fishes and crustaceans

Rates of predation on 0-group plaice, Pleuronectes platessa . in aquaria were compared under four different combinations of conditions to test the hypothesis that the presence of sand in which they may bury affords a refuge from predators. The effect of light and darkness on predation rate was also examined, Two crustaceans, the shrimp, Crangon crangon , and the portunid crab, Liocarcinus holatus , and two fishes, cod, Gadus morhua , and pollack, Pollachius pollachius , were used as predators. Predaton rates were significantly higher in the dark for all predators except pollack. Predation rates in the absence of sand were signifcantly greater only for pollack. The results suggest that predation rates on plaice during their juvenile nursery stage on sandy beaches will be significantly greater during darkness than during the day. Burying in sand appears to provide only a partial refuge from predation, perhaps because natural predators have evolved effective methods of foraging for buried prey.     

Diel vertical migrations of invertebrate predators (Leptodora kindtii ,Thermocyclops taihokuensis, andMesocyclops sp .) in a shallow, eutrophic lake

We studied vertical distribution patterns of three invertebrate predators – Leptodora kindtii, Mesocyclopssp., and Thermocyclops taihokuensis– in a shallow eutrophic lake, Lake Suwa , Japan. From June to October in 2000 and 2001, we collected samples in the lake center in order to examine the vertical distribution patterns and the densities of the predators in the water column during the day (0900) and at night (2330). We also examined phototactic behavior of Leptodora in the laboratory. The three invertebrate predators showed clear migration patterns. Leptodora and Thermocyclops displayed a typical migration, avoiding the surface and maintaining a high abundance in deeper water during the day, and being distributed uniformly during the night. Mesocyclops, on the other hand, showed no clear vertical distribution pattern in the water column. However, Mesocyclops showed higher densities in the water column during the night than during the day. It suggests that they stayed just above the bottom during the day and migrated upward during the night. Leptodora also showed such a density difference between day and night. In the laboratory, Leptodora showed strong negative phototactic behavior. The observed density changes between day and night in Leptodora and Mesocyclops suggests the possible underestimation of their population density by usual sampling methods, and thus the impact of predation on populations of prey zooplankton species may also be underestimated in shallow water bodies.     

Mortality, behavior and the effects of predators on the intertidal distribution of littorinid gastropods

Predators can affect the vertical distribution of mobile intertidal invertebrates in two ways: they can (1) cause greater mortality of prey at certain intertidal levels, and (2) induce prey to seek safer intertidal areas. In this study, we investigate whether low-intertidal and subtidal predators affect the intertidal distribution of two congeneric species of small herbivorous gastropods of northeastern Pacific shores, Littorina sitkana Philippi 1846, and L. scutulata Gould 1849. In particular, we tested the hypothesis that predators affect the distribution of these snails by inducing them to seek higher and safer intertidal areas. On a wave-sheltered shore in Barkley Sound, British Columbia, L. sitkana and L. scutulata were both killed by predatory crabs (e.g., Cancer productus) more frequently when tethered near the lower limit of their intertidal distribution ( approximately 1 m) than when tethered where they were most common ( approximately 2.5 m), suggesting that high mortality rates are partly responsible for the lower-limit of these snails' intertidal distribution. However, two field mark-recapture experiments indicated that the snails' behavioral response to predation risk also influences their distribution. In the first experiment, snails from the 2.5-m level (low risk) transplanted to the 1.0-m level (high risk) displayed a strong and consistent tendency to move shoreward, especially L. sitkana, some traveling 10-15 m in 2-3 days to regain their original level. These shoreward movements were especially precise in the northern part of the study area, where predation rates on tethered snails were greatest. Furthermore, larger more vulnerable snails were more strongly oriented shoreward than smaller individuals, indicating that antipredator behavior might also contribute to intertidal size gradients in these species. In the second mark-recapture experiment, we manipulated predation risk using small cages and found that snails exposed to the odors of C. productus crabs foraging on conspecific and heterospecific snails displayed more precise (L. sitkana and L. scutulata) and longer (L. sitkana) shoreward movements than snails held in control conditions. These results provide the first experimental evidence that antipredator behavior may contribute to the intertidal distribution patterns of littorinids.     

Predation on 0-group and older year classes of the bivalve Macoma balthica: interaction of size selection and intertidal distribution of epibenthic predators

The bivalve Macoma balthica is a common species in the Wadden Sea and North Sea. Juveniles temporarily use nurseries in the high intertidal. To explain this nursery use, predation pressure was examined for both juvenile and adult Macoma at low and high tidal flats. The study was carried out in the eastern Dutch Wadden Sea.Shrimps Crangon crangon, adult crabs Carcinus maenas, gobies Pomatoschistus and juvenile flatfish were more abundant and larger on low than on high tidal flats, but 0-group Carcinus was more abundant on the high tidal flats. Crangon and 0-group Carcinus stomachs frequently contained Macoma remains. These predators selectively preyed on small 0-group Macoma, both in the field and in laboratory experiments. The effect of predation by epibenthic animals and birds, on the low and high tidal flats, was examined in exclosure experiments (2 mm mesh). There was no effect of epibenthos exclosure on adult Macoma. For 0-group Macoma, densities were higher in exclosures than in the controls where predators had normal access. The density reduction by epibenthic predators was much larger in the low than in the high intertidal. We found no effect of bird predation on densities of 0- and 1+group Macoma.Thus, 0-group Macoma is under high predation pressure by epibenthos in the low intertidal, especially by shrimps, while they are relatively safe in the high intertidal. However, most of the shellfish outgrow their epibenthic predators during their first summer. Therefore, it becomes safe for the bivalves to redistribute to locations where epibenthic predators are abundant, during their first winter. On the other hand, it did not become clear from this study why many of the larger Macoma leave the high intertidal. Concluding, the nursery use of Macoma-spat in the high intertidal is probably, at least partly, an adaptation to avoid epibenthic predation.     

Effects of Tidal and Diel Cycles on Dugong Habitat Use

ABSTRACT Quantifying the factors influencing behaviors of aquatic mammalian grazers may enhance the generic understanding of grazer ecology. We investigated diel and tidal patterns in movements of the dugong (Dugong dugon) by Global Positioning System—tracking 12 animals in 5 inshore—intertidal and 3 offshore—subtidal habitats along the coast of Queensland, Australia. We examined effects of tide height and time of day on the dugong's distance from 1) the nearest coast, 2) water >3 m deep, 3) actual water depth (bathymetry + tide ht) experienced, and 4) distribution of the directions of movements. Both tidal and diel cycles influenced dugong movement. Tracked dugongs tended to be closer to shore at high tide than at low tide and closer to shore at night than during the day. Onshore movement was more prevalent on incoming tides and in the afternoon and evening. Offshore movement was more prevalent on outgoing tides and from midnight through the morning until midday. Tidal and diel variation in water depths used by the inshore—intertidal dugongs was small, but probably underestimated, hidden by a sampling bias in the telemetry equipment. Onshore movement at high tide allowed dugongs to exploit intertidal seagrass beds. Dugongs are closer to shore in afternoons and evenings than in mornings. This behavior may be related to the avoidance of predators or watercraft. Our findings can be used to predict spatial patterns of dugongs within areas of conservation management significance and to assess, avoid, and mitigate adverse effects of anthropogenic disturbance.     

Horizontal distribution of pelagic zooplankton in relation to predation gradients

An investigation was conducted examining the horizontal and vertical distribution of zooplankton in Lake Miramar, a southern California reservoir. Daphnia and Mesocyclops populations were most abundant offshore and in deeper water during the day but appeared to move toward shore and upward at night. The results of inshore zooplankton sampling provided no evidence chat the diel horizontal migration pattern was a result of sampler avoidance by zooplankton. Inshore-offshore differences in Daphnia and Mesocyclops abundance and diel migrations were reduced during winter and early spring. Rotifer zooplankters exhibited less seasonal variation in their horizontal distributions than did the large crustacean zooplankters at all times of the year. It is hypothesized that the spatial distribution of zooplankton is related to predation gradients in Lake Miramar. The dominant planktivore in the reservoir, young-of-the-year Micropterus salmoides. was abundant from late May through December and much less so from January to early May. They were largely restricted to the littoral zone and this produced horizontal gradients of planktivory which varied in strength seasonally and from day lo night. It appears that crustacean zooplankton in Lake Miramar avoid areas with abundant planktivores during the day but migrate into these areas at night when the intensity of planktivory is reduced. Rotifers exhibit less horizontal heterogeneity and no significant diel migrations, which is attributed to the reduced risk of predation that rotifers experience relative to crustacean zooplankters. A graphical model is proposed to integrate our understanding of diel vertical and horizontal migrations of zooplankton. In this model, gradients of predation are completely vertical in offshore areas and strongly horizontal in near shore areas. Gradients of food availability are roughly similar to those of predation intensity. Plankiers respond to these gradients by migrating in a path parallel to gradients of predation at dawn and parallel to gradients of food availability after dark.     

Temporal variation in habitat selection breaks the catch‐22 of spatially contrasting predation risk from multiple predators

Predator avoidance depends on prey being able to discern how risk varies in space and time, but this is made considerably more complicated if risk is simultaneously present from multiple predators. This is the situation for an increasing number of mammalian prey species, as large carnivores recover or are reintroduced in ecosystems on several continents. Roe deer Capreolus capreolus in southern Norway illustrate a case in which prey face two predators with contrasting patterns of predation risk. They face a catch‐22 situation: spatially avoiding the risk from one predator (lynx Lynx lynx in dense habitat) implies exposure to the other (hunters in open habitat). Using GPS‐data from 29 roe deer, we tested for daily and seasonal variation in roe deer selection for habitat with respect to the habitats’ year‐round average risk level. Generally, roe deer altered their habitat selection between night and day in a pattern consistent with being able to avoid predicted risk from the nocturnal lynx during night and predicted risk from human hunters during day. However, seasonal variation in habitat selection only partially corresponded with the predicted seasonal variation in risk. Whereas roe deer avoided areas with high risk from hunters more strongly during the hunting season than in other seasons, there was a lack of selection towards areas and time periods lowering the risk of lynx predation during winter. It seems likely that the risk of starvation and thermal stress constrain roe deer habitat selection during this energetically challenging season with cold temperatures, snow cover and limited natural forage. The habitat selection pattern of roe deer fits thus only partly with the two contrasting risk gradients they face. Adjusting risk‐avoidance behavior temporally can be an adaptive response in the case of several predators whose predation patterns differ in space and time.     

Dome-shaped functional response induced by nutrient imbalance of the prey

Nutritional ecological theory predicts that predators should adjust prey capture and consumption rates depending on the prey's nutritional composition. This would affect the predator's functional response, at least at high prey densities, i.e. near predator satiation. Using a simple fruitfly-wolf spider laboratory system in Petri dishes, we found that functional responses changed from day to day over a 7 day period. After 1 to 2 days of feeding, dome-shaped functional responses (i.e. reduced predation at highest prey densities) appeared in spiders fed nutritionally imbalanced prey, compared with steadily increasing or asymptotic functional responses with nutritionally near-optimal prey. Later again (days 5-7), the difference disappeared as the level of the functional response was reduced in both treatments. Experiments with adult females in spring and subadult spiders in autumn revealed opposite patterns: a dome-shaped response with high-lipid prey for reproductive females, for which protein-rich prey are optimal, and a dome-shaped (or simply reduced) response with high-protein prey for pre-winter subadults, for which high-lipid flies are the optimal prey. Our results have implications for predation theory and models of biological control that have, so far, neglected nutritional aspects; in particular, the dynamic nutritional state of the predators should be incorporated.     

Piscine predation on juvenile fishes on a Scottish sandy beach

Predation by larger fishes is a major cause of mortality for the populations of juvenile fishes on a sandy beach on the west coast of Scotland. Predation was concentrated on the most numerous species (0-group Pleuronectes platessa ) in June but with the decline in numbers and growth in size of this species, the fish predators had changed their diet in August to feed principally on small sandeels (Ammodytidae). Six major predatory species had taken 95 per cent of all fish found in predators' stomachs and could be roughly divided into three categories according to their piscivorous tendencies and their abundance. (1) Common species in which fishes formed the main item of their diet ( Gadus morhua ). (2) Species in which fishes formed only a relatively minor constituent of the diet but which, by virtue of their abundance, potentially had a significant predatory impact (I-group P. platessa ). (3) less abundant species whose diet consisted principally of fishes. The intensity of predation was generally greatest at night for G. morhua but during the day for I-group P. platessa. The distributions and movements of the dominant prey species can be interpreted in the light of their predator-prey relationships. The most common benthic species, 0-group P. platessa , move off -and onshore with the ebb and flow of the tides and concentrate in shallow water at night. Such movements would have the effect of maintaining the majority of the population in depths where predation pressure from nocturnally onshore-migrating gadoids and other predators is likely to be minimized.     

Diel vertical and horizontal distribution of crustacean zooplankton and young of the year fish in a sub-alpine lake: an approach based on high frequency sampling

Understanding the spatial dynamics of predators and their preyis one of the most important goals in aquatic ecology. We studiedspatial and temporal onshore–offshore distribution patternsin young of the year (YOY) Eurasian perch (Perca fluviatilis)and crustacean zooplankton (Daphnia hyalina, Cyclops prealpinus)along a transect in Lake Annecy (France). Our study representsa first attempt at coupling hydroacoustic fish survey and highfrequency zooplankton recording to assess simultaneously thelarge-scale distribution patterns of YOY fish and their zooplanktonprey over a diel cycle (day, dusk and night sampling). We hypothesizedthat the spatial distribution of zooplankton could be shapedby both anti-predator behaviour (horizontal and vertical migrations)and predation losses. Fish biomass, size structure and dietwere assessed from split-beam echosounding and net trawlingsamples, whereas crustacean abundances were estimated with asmall modified Longhurst–Hardy continuous plankton recorder.We evaluated the diel changes in the spatial distribution patternsof fish and zooplankton and determined the overlap between theirdistributions. Fish biomass was dominated by YOY perch in upperwarmer layers and salmonids (Coregonus lavaretus and Salvelinusalpinus) in the colder and oxygenated deep layers. YOY perchwere aggregated in dense schools in the epilimnion during theday and dispersed at night. Fish biomass was distributed alonga strong increasing onshore–offshore gradient at night,whereas crustacean prey showed a decreasing gradient. This onshore–offshorenegative gradient in crustacean distribution, expressed on ashorter scale during the day, shifted toward the surface watersat night. A distinct kinetic of diel vertical migration (DVM)patterns was exhibited by daphnid and cyclopoid populationsand resulted in distinct vulnerability to perch predation. Spatio-temporaldistribution of crustaceans in Lake Annecy during the diel cyclestudy was probably shaped both by predation loss to YOY perchand by anti-predator behaviour (DVM, DHM) by zooplankton. Theimplications for fine-scale studies of fish-zooplankton interactionsare discussed.     

Factors influencing the characteristics of fish assemblages in a large subtropical marine embayment

Mean number of species and density of fishes in nearshore shallow waters of Shark Bay, a large subtropical embayment, were c . seven and 19.5 times greater in seagrass than over bare sand, where protection from predators and the abundance of potential invertebrate prey were less. The number of fish species and density of fishes over bare sand were lower in nearshore than offshore waters, where there was a greater amount of organic material and thus presumably a greater density of benthic macroinvertebrate prey. Species composition in vegetated and unvegetated habitats differed markedly, with species such as Monacanthus chinensis, Apogon rueppellii and Pelates quadrilineatus being largely confined to seagrass, whereas others such as Pseudorhombus jenynsii, Torquigener whitleyi and Engyprosopon grandisquama were found predominantly or exclusively over bare sand. The ichthyofauna in beds of Posidonia australis , in which the canopy is uniformly dense, differed in composition and comprised a greater number of species and density of fishes than that in Amphibolis antarctica , in which an open space is present beneath the terminal clusters of relatively short leaves. Species composition in the beds of both of these seagrass species underwent well defined cyclical changes, caused by out-of-phase sequential changes in the densities of certain species. Such changes were less common over bare sand, where the ichthyofaunal composition was more variable. The number of species and density of fishes over bare sand were greater at night than during the day, reflecting, in part, a tendency for species such as A. rueppellii to move into unvegetated areas to feed at night, when the likelihood of predation by visual predators would be reduced. Within Shark Bay, ichthyofaunal composition is influenced most by habitat type (vegetated v . unvegetated), followed in general by water depth and then region in the bay and time of year.     

Activity patterns of insectivorous bats and birds in northern Scandinavia (69° N), during continuous midsummer daylight

Previous studies suggest that many species of insectivorous bats are nocturnal, despite the relatively low availability of their insect prey at night, because of the risk of predation by diurnal predatory birds. We hypothesised that if this was the case bats living above the arctic circle would alter their feeding behaviour during midsummer because there would no longer be any benefit to restricting their activity to the period when their prey are least abundant. Alternatively, if bats were more influenced by competition from aerial insectivorous birds they would continue to feed at 'night' to avoid such competition. In northern Norway (69°  N), during continuous midsummer daylight, insectivorous sand martins ( Riparia riparia ) concentrated their aerial feeding activity when aerial insects were most abundant. The birds stopped feeding between 23:00 and 07:00 when aerial insects were least abundant. In contrast, northern bats ( Eptesicus nilssonii ), fed mostly between 22:00 and 02:00, coinciding with the lowest aerial insect availability, and with the period when light levels were lowest (ca 1000 lux). Bat activity patterns were closest to those predicted by the avian competition hypothesis. The low densities of both sand martins and Northern bats in the study area, however, were less consistent with this hypothesis. Possibly populations of both species were higher historically and the observed patterns reflected historical competition. Bat activity was most closely correlated to ambient light levels. This raised two alternative explanations that we could not eliminate. Perhaps there was differential predation risk, between the brightest and darkest parts of the day, because the visual capacities of falcons are strongly dependent on luminance. Alternatively the bats may have been entrained to emerge at given light levels by their behaviour at other times of year.