The influence of predator regime on the behaviour and mortality of a freshwater amphipod, <Emphasis Type="Italic">Gammarus pulex</Emphasis>

In species with restricted dispersal, traits may become genetically fixed leading to local adaptations. Therefore, predator avoidance in a prey species may differ between populations experiencing different predator regimes, but also between sexes within a population due to different vulnerability to predators. In this study we used male and female Gammarus pulex from two different predator regimes: fishless ponds, where invertebrates are the dominant predators and ponds with predatory fish. In the laboratory we examined refuge use, mortality, leaf decomposition rate and pair-formation in G. pulex when exposed to predator cues from either invertebrate predators or fish. Individuals from fish ponds spent more time in refuge and had a higher mortality than those from fishless ponds independent of predator cues. There was no effect of pond predator regime or predator cues on leaf decomposition rates. Further, fewer individuals formed pairs in G. pulex from fish ponds than from fishless ponds. Male G. pulex had a higher mortality and a higher decomposition rate than females independent of predator cues. However, there was no difference in refuge use between sexes. Our study shows that there are general differences in behaviour traits, both between predator regimes and sexes in G. pulex.     

The influence of nutritional state on shoal choice in zebrafish, Danio rerio

We investigated the mechanisms and functions of shoal choice in relation to nutritional state in the zebrafish. Single fish that had been well fed or food deprived for 2 days were presented with a choice between two stimulus shoals. Food-deprived test fish showed a significant preference for well-fed stimulus fish over food-deprived ones whereas well-fed test fish did not exhibit any significant preference. Subsequent experiments showed that food-deprived test fish had a significantly higher foraging success in shoals consisting of well-fed individuals than in ones that comprised food-deprived fish. No difference in the locomotory behaviour of food-deprived and well-fed stimulus fish was found with respect to the proportion of time spent swimming (as opposed to being motionless), the proportion of time spent in the upper part of the test tank and the number of sharp turns. However, body weight, stomach width (measured directly behind the pectoral girdle) and ventro-dorsal height significantly decreased over a 48-h food-deprivation period. The potential use of the latter factors for the recognition of food-deprived individuals is discussed. Copyright 1999 The Association for the Study of Animal Behaviour.     

Training fish for restocking: refuge and predator training in the hatchery has limited benefits for a marine fish

Large numbers of hatchery-reared fish are released in stocking programmes; however, success is limited by high mortality. Predation is seen as the main cause of deaths but might be reduced by training fish before release to avoid predators and/or use refuge. In this study on a potential restocking species, yellowfin bream Acanthopagrus australis, the effects of predator training and refuge on the behaviour of fish in the hatchery were tested. In the first experiment, juvenile bream were exposed to predatory mangrove jack (Lutjanus argentimaculatus) fed exclusively on bream flesh while housed in tanks with and without refuge. Predator training altered fish behaviour when fish were re-exposed to predators, but the effects were subtle and varied between groups of fish. In contrast, refuge created strong and consistent changes in behaviour, significantly slowing down the amount of time that fish took to consume food. A second experiment focused on the effects of refuge. Bream were trained to use artificial seagrass or house bricks as refuge and then exposed to mangrove jacks in a laboratory predation experiment. When refuge was available, fish significantly slowed down their feeding rate. There was a small, transient increase in survival for fish given seagrass refuges, but this was irrespective of whether the bream were trained to use refuge. The results of this study indicate that the use of refuge may be innate and the benefits of refuge may be available to naive hatchery-reared fish or fish trained to use refuge shortly before release. This suggests that there is potential to improve post-release survival of fingerlings without time-consuming and expensive hatchery training.     

Biomanipulation: the next phase — making it stable

Review of the literature on biomanipulation shows that fish manipulations have occurred through the following methods: piscivore addition; piscivore catch restriction; habitat enhancement; piscivore removal; planktivore exclusion; planktivore removal by selective catch, lake emptying, fish poisons, fish diseases, winterkill, summerkill; habitat expansion or contraction; planktivore addition; and natural events. The methods can be classified as deliberate, inadvertent, or natural, all of which have successful and unsuccessful examples. However, the problem of perpetuation of successful results remains unsolved. It is proposed that this may be resolved through use of a refuge or refuges from among the following: low light intensity refuge; low temperature refuge; low dissolved oxygen refuge; physical concealment refuge; visual clutter refuge; behavior modification refuge; and predator inefficiency refuge. Perhaps through use of such mechanisms large herbivorous zooplankters can continue to exist in lakes. Examples are given. Contribution no. 383 from the Limnological Research center.     

Effects of nutritional state on the shoaling tendency of banded killifish, Fundulus diaphanus, in the field

Although group living can confer benefits to individuals in terms of reduced predation risk and enhanced foraging success, it may also be associated with costs, such as increased competition for food. The nutritional state of an individual could therefore affect its readiness to join or remain in groups. We investigated the shoaling behaviour of banded killifish by following marked individuals, whose nutritional state had been experimentally manipulated, and recording their shoaling behaviour in a field enclosure containing unmarked conspecifics. Overall, food-deprived fish spent more time alone, and therefore less time shoaling, than well-fed individuals. When shoaling, however, food-deprived fish were not found in smaller shoals than well-fed conspecifics. Furthermore, they did not show a greater latency to shoal initially. Having joined a shoal, however, food-deprived fish left shoals more frequently to be alone than fed fish. Rates of change in the membership size of shoals occupied by either well-fed or food-deprived fish did not differ. We conclude that nutritional state seems to affect an individual's decision to continue shoaling once an association has been made. This study is the first to investigate experimentally state-dependent changes in the size of social groups in fish under field conditions. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Feeding and Refuge Use by Small Fish in the Presence of Cyanobacteria Blooms

We studied the effect of cyanobacteria on foraging and refuge use in small fish. We measured pike larval feeding in the presence of cyanobacteria by counting leftover prey. Our results showed that feeding by pike larvae on zooplankton prey decreased significantly in the presence of non-toxic cyanobacteria. The behaviour can be due to lowered vision caused by turbidity or clogging of the gills. Further, we tested whether the three-spined stickleback use toxic cyanobacteria as a refuge against predators in a choice experiment. The choice experiment was performed in a Y-maze fluviarum, where the fish could select between two different environments. Our results support the refuge use hypothesis because the three-spined stickleback clearly preferred toxic cyanobacteria to the chemical predator signal. To conclude, cyanobacteria decrease feeding rates in fish larvae, but may function as important refuge for e.g. sticklebacks, during predation pressure in pelagic algal blooms.     

Genetic mixed-stock analysis of Atlantic sturgeon Acipenser oxyrinchus oxyrinchus in a heavily exploited marine habitat indicates the need for routine genetic monitoring

Although a previous genetic mixed-stock analysis (gMSA) conducted in the early 1990s showed that marine-captured New York Bight Atlantic sturgeon Acipenser oxyrinchus oxyrinchus almost exclusively originated from the Hudson River, fish from southern U.S. rivers were well represented within this contemporary sample (n = 364 fish), at least during the autumn. Widely distributed spawning stocks are therefore exposed to heavy fishing activity and habitat degradation in this relatively small area, illustrating the need for spatial management across multiple management jurisdictions and routine gMSA to account for temporal change.     

Disturbance chemical cues determine changes in spatial occupation by the convict cichlid Archocentrus nigrofasciatus

I studied the effect of disturbance chemical cues on fish that make trade-offs between foraging in an open area and remaining in a safe refuge. I used convict cichlids Archocentrus nigrofasciatus that were either visually exposed to a predator (n = 8) or exposed to water conditioned by chemical cues from disturbed conspecifics (n = 8). Fish visually exposed to a predator decreased their ingestion rate and spent more time in the refuge than in the foraging area, while fish receiving water from frightened conspecifics did not alter their ingestion rate or time spent in the refuge and foraging site, but increased their spatial occupation (i.e., motion). These results suggest that convict cichlids recognized the predator by visual cues. Moreover, disturbance cues are a form of threatening public information that may increase fish spatial occupation due to increased exploring behaviour; but is not sufficiently alarming to alter feeding or increase refuge use.     

Hunger, light level and body size affect refuge use by post-settlement lingcod Ophiodon elongatus

Using experimental methods, body size, nutritional state (hunger) and light level were investigated as possible determinants of the extent to which juvenile lingcod Ophiodon elongatus used structural refuges. In the laboratory, older, larger individuals (134–162 mm total length, L _T) exhibited a higher propensity for structure use than did smaller juvenile fish (78–103 mm L _T). In fish of both body sizes, increasing hunger levels caused fish to emerge more readily from refuges than satiated individuals, with this effect most prominent for smaller fish. Diel patterns of ambient illumination were simulated in the laboratory, and refuge use by large individuals decreased significantly in response to lower nocturnal light levels, a pattern that was not observed in the smaller size class. These results indicate an ontogenetic shift in refuge use. During the summer, as post-settlement fish grew larger, they tended to increase affinity for structurally complex habitat, and also showed divergent patterns of behaviour in relation to hunger and light level.     

Dispersal Behavior, Boldness, and the Link to Invasiveness: A Comparison of Four Gambusia Species

Dispersal is a key element of a species' invasiveness. Although considerable work has addressed how dispersal influences the pattern of spatial spread of invading organisms, few studies investigate whether invasive species are in fact better dispersers than either the species they displace or less successful invaders. Recent work suggests that variation in dispersal may be due to variation in an underlying behavioral trait, boldness. Our study examined the link between dispersal, boldness, and invasiveness by comparing the dispersal characteristics and refuge use of two invasive Gambusia species to two congeners in experimental streams. The streams consisted of a series of pools (no flow) connected to a flowing channel. For each species, small groups of females were released at the middle pool, and their movement and activity were recorded over a 1-h period. We found invasive Gambusia to be more likely to disperse out of the introductory pool, to disperse sooner, to travel a greater distance in the artificial streams, and thus to exhibit greater dispersal tendencies than their close relatives. Among the invasives, Gambusia affinis had a greater dispersal tendency than G. holbrooki. We suspect this result indicates variation in the contribution of dispersal to the relative invasiveness of these species. Certain dispersal measures were correlated to time spent out of refuge, although invasive Gambusia and their relatives did not differ in the predicted manner. These results argue for the greater incorporation of experimental approaches and analyses of behavioral mechanisms in the study of invasive species.     

Movement patterns,home range size and habitat utilization of the bluespine unicornfish, <Emphasis Type="BoldItalic">Naso unicornis</Emphasis> (Acanthuridae) in a Hawaiian marine reserve

Synopsis We quantified bluespine unicornfish, Naso unicornis, movement patterns, home range size and habitat preferences in a small Hawaiian marine reserve. Bluespine unicornfish were site-attached to home ranges situated within the reserve boundaries and their movements were aligned with topographic features. Two different diel movement patterns (‘commuting’ and ‘foraying’) were observed. Commuters made crepuscular migrations of several hundred meters between daytime foraging areas and nighttime refuge holes. Foraying fish did not partake in crepuscular migrations and utilized refuge holes both day and night. Two bluespine unicornfish were also nocturnally active. There was little direct evidence of dispersal from the reserve but differences in bluespine unicornfish abundance and size among reef habitat zones were consistent with ontogenetic habitat shifts. The influence of habitat topography on bluespine unicornfish movements suggests that gross habitat characteristics could be used to predict reef fish movements. This could provide a simple method for setting marine reserve boundaries at sites for which empirical fish movement data are unavailable.     

Refuge use by fish as a function of body weight changes

Refuge use provides a good model for the study of trade-offs between the benefits of predator avoidance and the costs of lost feeding opportunities. We manipulated the latter costs by subjecting similar-sized three-spine sticklebacks to 2 days of food deprivation followed by a 2-day re-feeding period and recorded associated changes in body weight and refuge use. Food deprivation resulted in a decrease and re-feeding in an increase in the duration of refuge use by fish. Emergence times of fish from the refuge were extremely variable (with a ratio of 1:127 between the shortest and the longest ones) but individual ranks were highly consistent between different days of testing, suggesting that emergence times were individually characteristic. Percentage weight change of fish in response to the experimental treatments also showed a high level of inter-individual variation ranging from 0–17%. A significant positive correlation was found between the percentage weight lost and the percentage decrease in emergence time from a refuge after food deprivation and similarly between the percentage weight gained and the percentage increase in refuge use after re-feeding. The relationship between energy turnover and behavioural strategies is discussed. Received: 13 December 1998 / Received in revised form: 2 May 1999 / Accepted: 7 June 1999     

Intraspecific variation in behaviour: effects of evolutionary history, ontogenetic experience and sex

Geographical variation in behaviour within species is common. However, how behavioural plasticity varies between and within locally adapted populations is less studied. Here, we studied behavioural plasticity induced by perceived predation risk and food availability in pond (low predation - high competition) vs. coastal marine (high predation - low competition) nine-spined sticklebacks (Pungitius pungitius) reared in a common garden experiment. Pond sticklebacks were more active feeders, more risk-taking, aggressive and explorative than marine sticklebacks. Perceived predation risk decreased aggression and risk-taking of all fish. Food restriction increased feeding activity and risk-taking. Pond sticklebacks became more risk-taking than marine sticklebacks under food shortage, whereas well-fed fish behaved similarly. Among poorly fed fish, males showed higher drive to feed, whereas among well-fed fish, females did. Apart from showing how evolutionary history, ontogenetic experience and sex influence behaviour, the results provide evidence for habitat-dependent expression of adaptive phenotypic plasticity.     

Diffusion models for population dynamics incorporating individual behavior at boundaries: applications to refuge design.

We construct models for dispersal of a population which incorporate the response of individuals to interfaces between habitat types. The models are based on random walks where there may be a bias in the direction an individual moves when it encounters an interface. This sort of dispersal process is called skew Brownian motion. Our models take the form of diffusion equations with matching conditions across the interface between regions for population densities and fluxes. We combine the dispersal models with linear population growth models which assume that the population growth rate differs between regions of different habitat types. We use those models to study issues of refuge design. We specifically consider how the effectiveness of buffer zones depends on their size, quality, and the population's response to the interface between the buffer zone and the refuge.     

The relationship between foraging and shoal position in a mixed shoal of roach (Rutilus rutilus) and chub (Leuciscus cephalus): a field study

Feeding rates of mixed shoals of juvenile roach and chub were observed in a shallow stream near Cambridge (UK). Roach at the front of the shoal had significantly higher feeding rates than roach at the back and than chub in either front or back positions. Position in the shoal also had a significant effect on the kind of food consumed, with front roach feeding more on plankton and back roach more on bottom food. Altogether 36 fish from the stream were caught and marked. Half of these were deprived of food and the other half well-fed for 3 days in captivity. After release 36% of them joined their old shoal again. Individuals from the starved group occupied front positions significantly more often than well-fed fish, but after 2 days this difference disappeared.     

Individual behaviour and resource use of thermally stressed brook trout Salvelinus fontinalis portend the conservation potential of thermal refugia

Individual aggression and thermal refuge use were monitored in brook trout Salvelinus fontinalis in a controlled laboratory to determine how fish size and personality influence time spent in forage and thermal habitat patches during periods of thermal stress. On average, larger and more exploratory fish initiated more aggressive interactions and across all fish there was decreased aggression at warmer temperatures. Individual personality did not explain changes in aggression or habitat use with increased temperature; however, larger individuals initiated comparatively fewer aggressive interactions at warmer temperatures. Occupancy of forage patches generally declined as ambient stream temperatures approached critical maximum and fish increased thermal refuge use, with a steeper decline in forage patch occupancy observed in larger fish. These findings suggest that larger individuals may be more vulnerable to stream temperature rise. Importantly, even at thermally stressful temperatures, all fish periodically left the thermal refuge to forage. This indicates that the success of refugia at increasing population survival during periods of stream temperature rise may depend on the location of thermal refugia relative to forage locations within the larger habitat mosaic. These results provide insights into the potential for thermal refugia to improve population survival and can be used to inform predictions of population vulnerability to climate change.     

On the Size of African Riverine Fish Faunas

SYNOPSIS. An apparent anomaly exists in the size of riverinefish faunas of the Nile and Zaire; the Nile while considerablylonger, appears depauperate when compared with the Zaire. Thisparadox is explained when discharge, not length, is used asa measure of river size. Indeed, the number of freshwater fishspecies in African rivers is more closely related to dischargethan to length or catchment area. Discharge is directly proportionalto terrestrial productivity of a river basin, which in turnaffects total biomass of fish and number of species. Changesin the size of rivers during the geologic past affected theircapacity to sustain diverse fish faunas. Rivers flowing throughespecially arid lands during the late- Pleistocene were reducedin discharge, and concomitantly, fish faunas. Immigration offish from refuge rivers during the Holocene partially restoredthese diminished faunas. We propose that fish are more mobilethan they seem, and that the distinctiveness of riverine fishfaunas may be maintained by competitive pressure from establishedresidents, rather than by limited dispersal abilities of fish.Theories of the distribution of fish in Africa are considered,and we suggest that discharge as affected by climatic stabilityis largely responsible for the size of African riverine fishfaunas.     

Fleeing towards death – leech‐induced behavioural defences increase freshwater snail susceptibility to predatory fish

Prey species are often exposed to multiple predators, which presents several difficulties to prey species. This is especially true when the response to one predator influences the prey’s susceptibility to other predators. Predator‐induced defences have evolved in a wide range of prey species, and experiments involving predators with different hunting strategies allow researchers to evaluate how prey respond to multiple threats. Freshwater snails are known to respond to a variety of predators with both morphological and behavioural defences. Here we studied how freshwater snails Radix balthica responded behaviourally to fish and leech predators, both separately and together. Our aim was to explore whether conflicting predator‐induced responses existed and, if so, what effect they had on snail survival when both predatory fish and leeches were present. We found that although R. balthica increased refuge use when exposed to predatory fish, they decreased refuge use when exposed to predatory leeches. When both predators were present, snails showed a stronger response towards leech than fish and responded by leaving the refuge. This response made the snails more susceptible to fish predation, which increased snail mortality when exposed to both fish and leech compared to fish only. We show that predators that have a relatively low predation rate can substantially increase mortality rates by indirect effects. By forcing snails out of refuges such as rock and macrophyte habitats, leeches can indirectly increase predation from molluscivorous fish and may thus affect snail densities.     

Refuge use by fish as a function of body length-related metabolic expenditure and predation risks

Theory predicts that staying in a refuge has benefits in terms of predator avoidance and costs in terms of lost feeding opportunities. In this study, we investigated how the relative importance of these costs and benefits changes with increasing body length. This is of particular interest in animals such as fish, which show continuous growth throughout their lives. Our results suggest that larger fish are subject to lower predation risks and are less affected by food deprivation than small fish, with fish decreasing their responses to food-deprivation treatments more strongly with increasing body length than to predation treatments. This may explain our observation that large fish emerged later from a refuge than small ones and spent shorter times outside the refuge. The key role of differential responses to food deprivation was further illustrated by the finding that the relative weight loss of individual fish was strongly correlated with a reduction in hiding time even in the absence of body length differences. The importance of inter-individual differences in metabolic rates for the decision-making behaviour of animals is discussed.     

The Effect of Hunger on Shoal Choice in Golden Shiners (Pisces: Cyprinidae,Notemigonus crysoleucas)

When an animal has a choice of joining one group over another, its decision may depend on its relative vulnerabilities to predation and starvation. For example, a well-fed animal may choose a large group of individuals with body size matching its own because this gives good protection against predators, but a hungry animal may prefer smaller groups made up of smaller individuals because this decreases food competition. To test this idea, a choice between various shoals was given to golden shiners, Notemigonus crysoleucas, that were either well fed or deprived of food for 48 h. In a choice of 10 vs. 3 shoalmates, both well-fed and hungry shiners spent more time near the shoal of 10. In a choice of 20 vs. 3 shoalmates, both well-fed and hungry shiners again preferred the larger shoal, but in one replicate this preference was significantly weaker in the hungry fish. This reduced preference did not appear to be an artefact of increased mobility by hungry fish searching for food. In a choice between shoals of small vs. large conspecifics, small well-fed shiners, small hungry shiners, and large well-fed shiners preferred shoalmates with body size matching their own, but large hungry shiners preferred smaller individuals. These results are consistent with the hypothesis that hungry fish sacrifice safety from predation in their shoaling behaviour (by avoiding larger groups to a certain extent and by risking the oddity effect) so as to decrease food competition.