What limits predator detection in blue tits (<Emphasis Type="Italic">Parus caeruleus</Emphasis>): posture,task or orientation?

To detect threats and reduce predation risk prey animals need to be alert. Early predator detection and rapid anti-predatory action increase the likelihood of survival. We investigated how foraging affects predator detection and time to take-off in blue tits (Parus caeruleus) by subjecting them to a simulated raptor attack. To investigate the impact of body posture we compared birds feeding head-down with birds feeding head-up, but could not find any effect of posture on either time to detection or time to take-off. To investigate the impact of orientation we compared birds having their side towards the attacking predator with birds having their back towards it. Predator detection, but not time to take-off, was delayed when the back was oriented towards the predator. We also investigated the impact of foraging task by comparing birds that were either not foraging, foraging on chopped mealworms, or foraging on whole ones. Foraging on chopped mealworms did not delay detection compared to nonforaging showing that foraging does not always restrict vigilance. However, detection was delayed more than 150% when the birds were foraging on whole, live mealworms, which apparently demanded much attention and handling skill. Time to take-off was affected by foraging task in the same way as detection was. We show that when studying foraging and vigilance one must include the difficulty of the foraging task and prey orientation.Communicated by P.A. Bednekoff     

Behavioral resource depression and decaying perceived risk of predation in two species of coexisting gerbils

Summary Behavioral resource depression occurs when the behavior of prey individuals changes in response to the presence of a predator, resulting in a reduction of the encounter rate of the predator with its prey. Here I present experimental evidence on the response of two species of gerbils (Gerbillus allenbyi and G. pyramidum) to the presence of barn owls. I conducted the experiments in a large aviary. Both gerbils responded to the presence of barn owl predators by foraging in fewer resource patches (seed trays) and by quitting foraged resource patches at a higher resource harvest rate (giving-up density of resource; GUD). This reduced the amount of time gerbils were exposed to owl predation, and hence the encounter rate of owls with gerbils, i.e., behavioral resource depression. Thus, the presence of owls imposes a foraging cost on gerbils due to risk of predation, and also on the owls themselves due to resource depression. I then examined how resource depression relaxed over time following exposure to owls. In the days following an encounter with the predator, the reduction in foraging activity for both gerbil species eased. Increasing numbers of trays were foraged each day, and GUDs in seed trays declined. The two gerbils differed in their rate of recovery, with G. pyramidum returning to prepredator levels of foraging after 1 or 2 nights and G. allenbyi taking 5 nights or longer. Interspecific differences in recovery rates may be based on differences between the species in vulnerability to predation and/or ability to detect the presence of predators. The differences in recovery rates may be due to optimal memory windows or decay rates, where differences between species are based on risk of predation or on how perceived risk changes with time since a predator was last encountered. Finally, differences between or among competitors in recovery from resource depression may provide foraging opportunities in time for the species which recover most quickly and may have implications for species coexistence.     

Predation risk affects courtship and attractiveness of competing threespine stickleback males

The effect of predation risk and male-male competition on male courtship behaviour and attractiveness to females was studied in the threespine stickleback (Gasterosteus aculeatus) by presenting dummy or live females to solitary and competing males under different predation risks. In the presence of a predator, males decreased courtship activity. Different courtship components were, however, adjusted to different extents and in opposing directions to predation risk, probably because the single components may have varied in riskiness. The presence of a competing male decreased overall courtship activity, but increased the frequency of zigzags, suggesting zigzagging to be a competitive strategy against other males. In the presence of a predator male courtship activity was not affected by a competitor. Female mate choice correlated with the males' previous frequency of zigzags towards a dummy female. However, when a live female paid attention to a male, the male decreased zigzagging and instead increased leading and fanning behaviours, probably trying to attract the female to the nest to mate. Predation risk affected the attractiveness of males as females reduced their attention to a male when he faced a predator and reduced his courtship activity. As females instead increased their attention to a competing male that had increased his courtship activity, due to decreased competition, males clearly are balancing mating opportunities against predator avoidance. When males vary in their susceptibility to predators, predation risk may thus affect mating success of competing males. Received: 31 January 1997 / Accepted after revision: 15 April 1997     

Adjustment of parental effort to manipulated foraging ability in a pelagic seabird,the thin-billed prion Pachyptila belcheri

An experiment was designed to examine in a long-lived seabird, the thin-billed prion (Pachyptila belcheri), how adults adjust their food provisioning strategy when their foraging abilities are reduced and when the chick's needs are increased. To reduce the foraging abilities of adults we impaired their flying ability by removing some flight feathers (handicapped), and to increase the food needs of the chick one parent was retained (single). Birds made either short foraging trips lasting 1–3 days, or long trips lasting 5–9 days. Control birds alternated long and short trips whereas single birds or handicapped birds made several successive short trips and thereafter a long trip. In each treatment, food loads tended to be heavier after long trips than after short trips, and single birds tended to bring heavier loads than control or handicapped birds. Birds in the three treatments lost similar amounts of mass after short trips and gained similar amounts of mass after long trips. However, the mass of handicapped birds declined through the experiment, while that of control and single birds remained stable. Although the proportion of chicks that died during the experiment was similar among the three treatments, the chicks fledged by a single bird were lighter than those in control nests. The results of the experiment suggest that thin-billed prions adjust their breeding effort differently to decreased flying ability or increased food demand by the chick. Single birds increase foraging effort without allowing their condition to deteriorate. Conversely, handicapped birds are unable to maintain their body condition while sustaining the chick at the same rate as control birds. It is suggested that in this long-lived seabird, adults probably adjust their breeding effort so that they do not incur the risk of an increased mortality, this risk being monitored by the body condition.     

Different antipredator behaviour in two anuran tadpoles: effects of predator diet

Recent investigations have indicated that animals are able to use chemical cues of predators to assess the magnitude of predation risk. One possible source of such cues is predator diet. Chemical cues may also be important in the development of antipredator behaviour, especially in animals that possess chemical alarm substances. Tadpoles of the common toad (Bufo bufo) are unpalatable to most vertebrate predators and have an alarm substance. Tadpoles of the common frog (Rana temporaria) lack both these characters. We experimentally studied how predator diet, previous experience of predators and body size affect antipredator behaviour in these two tadpole species. Late-instar larvae of the dragonfly Aeshna juncea were used as predators. The dragonfly larvae were fed a diet exclusively of insects, R. temporaria tadpoles or B. bufo tadpoles. R. temporaria tadpoles modified their behaviour according to the perceived predation risk. Depending on predator diet, the tadpoles responded with weak antipredatory behaviour (triggered by insect-fed predators) or strong behaviour (triggered by tadpole-fed predators) with distinct spatial avoidance and lowered activity level. The behaviour of B. bufo in predator diet treatments was indistinguishable from that in the control treatment. This lack of antipredator behaviour is probably related to the effective post-encounter defenses and more intense competitive regime experienced by B. bufo. The behaviour of both tadpole species was dependent on body size, but this was not related to predator treatments. Our results also indicate that antipredator behaviour is largely innate in tadpoles of both species and is not modified by a brief exposure to predators. Received: 22 August 1996 / Accepted after revision: 31 January 1997     

Autotomy shapes the trade-off between seeking cover and foraging in larval damselflies

Animals commonly choose between microhabitats that differ in foraging return and mortality hazard. I studied the influence of autotomy, the amputation of a body part, on the way larvae of the damselfly Lestes sponsa deal with the trade-off between foraging or seeking cover. Survival of Lestes larvae when confronted with the odonate predator Aeshna cyanea was higher in a complex than in a simple microhabitat, indicating that this more complex microhabitat was safer. Within the simple microhabitat, larvae without lamellae had a higher risk for mortality by predation than larvae with lamellae, showing a long-term cost of autotomy. When varying the foraging value (food present or absent) and predation risk (encaged predator or no predator) in the simple microhabitat, larvae with and without lamellae responded differentially to the imposed trade-off. All larvae spent more time in the simple microhabitat when food was present than when food was absent. Larvae without lamellae, however, only sporadically left the safe microhabitat, irrespective of the presence of the predator. In contrast, larvae with lamellae shifted more frequently towards the risky microhabitat than those without lamellae, and more often in the absence than in the presence of the predator. These decisions affected the foraging rates of the animals. I show for the first time that refuge use is higher after autotomy and that this is associated with the cost of reduced foraging success. The different microhabitat preferences for larvae with and without lamellae are consistent with their different vulnerabilities to predation and demonstrate the importance of intrinsic factors in establishing trade-offs. Received: 4 June 1999 / Received in revised form: 18 August 1999/ Accepted: 18 August 1999     

Energetic state and the performance of dawn chorus in silvereyes (Zosterops lateralis)

Stochastic dynamic programming (SDP) models predict that males singing to attract a mate should concentrate singing in what has been termed the dawn chorus. This is because male birds should have a variable surplus of fat in the morning that can be used to fuel singing, with the amount of fat available dependent upon such factors as his quality, foraging success and risk of predation. In this manner, the dawn chorus can act as an indicator of male quality in the context of female mate choice. We test a key prediction of SDP models of singing behaviour that males with greater fat levels should sing more. We conducted an experiment where we recorded the dawn chorus of male silvereyes (Zosterops lateralis) on three consecutive days. Each male received supplementary food on the second day, which enabled us to sample his dawn chorus before, during and after food supplementation. We also collected data on the effect of supplementary food on the body mass of silvereyes. As predicted by SDP models, we found that silvereyes sang for a greater proportion of the time after receiving supplementary food. Supplementary food also had a significant effect on the complexity of a male song, indicating that males not only increased the quantity of their song but also the quality of their song when they received extra food. As the provision of supplementary food significantly increased the mass of fed birds, our results support a causal link between male energy reserves and his ability to perform the dawn chorus.     

Timing of the evening emergence from day roosts of the grey-headed flying fox, Pteropus poliocephalus: the effects of predation risk, foraging needs, and social context

This study addresses the functional question of how variation in foraging strategy, predation risk, and social context influence the timing of the evening emergence from day roosts of the grey-headed flying fox, Pteropus poliocephalus. The onset of evening emergence was expected to vary according to the relative costs and benefits of emerging early and should, therefore, reflect an optimal trade-off between predation risks and foraging needs. The onset of the colony-wide emergence was closely correlated with the time of sunset and cloud cover. However, as expected, the onset of the colony-wide emergence was delayed when a diurnal avian predator was present, whereas the onset was advanced during lactation when presumably energetic demands are higher. The trade-off between predation risks and foraging needs was further reflected in the emergence times of individual bats: adult females emerged earlier when they had higher foraging needs as indicated by their body condition; young emerged later when they were smaller and likely to be more at risk from predation due to their less developed flying skills. However, the emergence time of adult males depended on their social status: smaller bachelor males emerged from the colony earlier than larger harem-holding males who guard their harems until the last female had left. Thus, whereas the colony-wide emergence time reflected the outcome of a trade-off between predation risks and general foraging needs, on an individual level, the outcome of this trade-off depended on sex, age, body condition, and structural size and was modified by social context.     

Feeding and aggressive behaviours in juvenile coho salmon (Oncorhynchus kisutch) under chemically-mediated risk of predation

Summary Juvenile coho salmon (Oncorhynchus kisutch) spend the first year of their lives in their natal streams, where they may often hold feeding territories. They also face significant risk of predation by birds and fish, and should alter their behaviour to reduce risk of mortality when these predators are present. Although there is laboratory evidence that coho react to predator visual stimuli, chemoreception of avian predator presence has not previously been reported. We tested the influence of chemical stimuli of common merganser (Mergus merganser), preying on juvenile coho, on two aspects of coho territorial behaviour, foraging and aggression, in flow-through aquaria. After a mixture of merganser- and coho-conditioned water was introduced into the system, juvenile coho significantly reduced their attack distance on drifting prey. The fish also significantly decreased their aggressive behaviour directed towards mirrors (total number of acts, intensity of acts and time spent) when the same odour was present. They did not change their behaviour in either experiment after control introductions of water treated with fish alone. These results are interpreted within the framework of a trade-off between juvenile growth and mortality.     

Shoal composition, body size and foraging in sticklebacks

Individuals which deviate from the majority in groups are likely to be most vulnerable to predation. This oddity effect, by definition, is frequency dependent, eventually fading at equal frequencies of the phenotypes in a group. It has been hypothesized that the increased predation risk of odd individuals may play an important role in the formation of phenotypically uniform shoals of fish. However, recent work has indicated that individuals may experience, or value, their predation hazard differently depending on their own size in relation to that of other group members: single large fish, but not small ones, appear concerned about their oddity in a shoal. Here I show that the apparent wariness of large fish is also expressed in a frequency-dependent manner, closely conforming to what is predicted if the oddity effect is responsible for their behavior. Using foraging activity of individuals as a means to evaluate their predation risk, I demonstrate with shoals comprising 12 threespine sticklebacks (Gasterosteus aculeatus) that large fish forage least actively when in a shoal consisting of 2 large and 10 small fish. An increase in the number of large fish to 4 among 8 small individuals clearly results in an increase in their foraging activity. However, having reached an equal frequency with small fish in a shoal, large fish do not seem to change their foraging activity much even when their number in a shoal increases further. In contrast, foraging activity of small sticklebacks remains fairly constant throughout the entire range of tested shoal compositions, providing further evidence that small and large fish respond to their oddity differently. Received: 12 February 1998 / Accepted after revision: 7 May 1998     

Stopover behaviour and departure decision of northern wheatears, Oenanthe oenanthe, facing different onward non-stop flight distances

On the small North Sea island Helgoland (54°11' N, 07°55' E) we studied the stopover ecology of two subspecies of northern wheatear, Oenanthe oenanthe, during spring migration. Birds heading for Scandinavia (O. o. oenanthe) face only short flights across an ecological barrier (50-500 km) whereas those originating from Greenland and Iceland (O. o. leucorhoa) have to cover between 1,000 and 2,500 km in the impending flight. Colour-ringed individuals showed that 90% of Scandinavian birds left on the day of ringing while 40% (males) and 30% (females) of Greenland/Icelandic birds stayed at least 1 night. The birds who remained were thus mostly O. o. leucorhoa. They often established desirable feeding territories on the beach and had a high rate of body mass increase (1.7 g/day). However, subspecies did not differ in habitat choice and in foraging effort, but O. o. leucorhoa had a higher success rate in pecking. Departure decisions were analysed by comparing (a) conditions on the day of ringing between departing and staying birds and (b) for birds staying between the day of departure and the preceding day. The factors that were probably important in the decision to depart differed between subspecies. In O. o. leucorhoa, few birds departed with bad or deteriorating weather conditions (tailwind component, cloud cover), whereas departures of O. o. oenanthe seemed to be little affected by those factors. A few O. o. oenanthe stayed early in the spring migration season and/or had low fat reserves. Interference during foraging seemed to play a role because both subspecies tended to leave when the densities of northern wheatears were high. Other factors related to refuelling conditions (food supply, foraging effort, predation risk) failed to show differences between staying and departing individuals. In summary, almost all Scandinavian birds departed quickly and irrespective of refuelling and weather conditions, whereas many (but not all) Greenland/Icelandic birds seemed to prepare for a long-distance flight and carefully adjusted departure to weather conditions. The observed differences in stopover behaviour and departure decisions in the two subspecies of northern wheatear indicate that the distance to the next stopover site or to the goal area has to be considered when applying optimal migration models.     

Threat-sensitive feeding strategy of immature sticklebacks (Gasterosteus aculeatus) in response to recent experimental infection with the cestode Schistocephalus solidus

Threat-sensitive decision-making might be changed in response to a parasitic infection that impairs future reproduction. Infected animals should take more risk to gain energy to speed up their growth to achieve early reproduction and/or to strengthen their immune response. To avoid correlational evidence, we experimentally infected and sham-infected randomly selected immature three-spined sticklebacks with the cestode Schistocephalus solidus. For 7 weeks we determined the threat-sensitive foraging decisions and growth of individual sticklebacks in the presence of a live pike (Esox lucius). The experimenters were blind with respect to the infection status of the fish. In contrast to previous studies, our recently infected fish should have been almost unconstrained by the parasite and thus have been able to adopt an appropriate life history strategy. We found a strong predator effect for both infected and uninfected fish: the sticklebacks’ risk-sensitive foraging strategy resulted in significantly reduced growth under predation risk. Infected fish did not grow significantly faster under predation risk than uninfected fish. Since infected fish consumed much less prey in the presence of the predator than did infected fish in its absence, they obviously did not use the opportunity to maximize their growth rate to reach reproduction before the parasite impairs it. Received: 21 June 1999 / Revised: 27 November 1999 / Accepted: 5 September 2000     

Behavioural responses to indirect and direct predator cues by a mammalian herbivore, the common brushtail possum

Cues for detecting and responding to perceived predation risk may be indirect, i.e., correlated with the probability of encountering a predator, or direct, i.e., produced by or related to the actual presence of a predator. Research shows, independently, both types of cues can influence anti-predator and foraging behaviours in prey species. However, since animals naturally encounter indirect and direct cues simultaneously, we were interested in quantifying their cumulative effect. Our aim was to evaluate food intake and behaviours (patch use, feeding (rate and time), vigilance) of a nocturnal mammalian herbivore to indirect (open vs. covered microhabitats; illumination) and direct (fox/owl odours) predator cues. We ran a preference trial with four paired treatments using a covered Safe food patch and an open Risk food patch, with one of four combinations of indirect and direct predator cues. Predation risk had a significant effect on both intake and behaviour (including feeding time, rate, and vigilance), but these effects differed depending on cues. No two combinations of cues produced exactly the same effects, illustrating the complexity of interactions that occur between cues. Covered patches were always perceived as less risky than open patches, but unexpectedly, open patches were perceived as riskier when dark rather than light. The strongest suite of (negative) responses to risk was associated with combined indirect and direct cues. These results highlight the importance of considering jointly, intake from a patch, intake rate, and behaviours, such as the proportion of time spent vigilant, when quantifying predation risk, rather than intake alone.     

The growth-predation risk trade-off under a growing gape-limited predation threat

Growth is a critical ecological trait because it can determine population demography, evolution, and community interactions. Predation risk frequently induces decreased foraging and slow growth in prey. However, such strategies may not always be favored when prey can outgrow a predator's hunting ability. At the same time, a growing gape-limited predator broadens its hunting ability through time by expanding its gape and thereby creates a moving size refuge for susceptible prey. Here, I explore the ramifications of growing gape-limited predators for adaptive prey growth. A discrete demographic model for optimal foraging/growth strategies was derived under the realistic scenario of gape-limited and gape-unconstrained predation threats. Analytic and numerical results demonstrate a novel fitness minimum just above the growth rate of the gape-limited predator. This local fitness minimum separates a slow growth strategy that forages infrequently and accumulates low but constant predation risk from a fast growth strategy that forages frequently and experiences a high early predation risk in return for lower future predation risk and enhanced fecundity. Slow strategies generally were advantageous in communities dominated by gape-unconstrained predators whereas fast strategies were advantageous in gape-limited predator communities. Results were sensitive to the assumed relationships between prey size and fecundity and between prey growth and predation risk. Predator growth increased the parameter space favoring fast prey strategies. The model makes the testable predictions that prey should not grow at the same rate as their gape-limited predator and generally should grow faster than the fastest growing gape-limited predator. By focusing on predator constraints on prey capture, these results integrate the ecological and evolutionary implications of prey growth in diverse predator communities and offer an explanation for empirical growth patterns previously viewed to be anomalies.     

Toward a predictive theory of risk effects: hypotheses for prey attributes and compensatory mortality

Risk effects, or the costs of antipredator behavior, can comprise a large proportion of the total effect of predators on their prey. While empirical studies are accumulating to demonstrate the importance of risk effects, there is no general theory that predicts the relative importance of risk effects and direct predation. Working toward this general theory, it has been shown that functional traits of predators (e.g., hunting modes) help to predict the importance of risk effects for ecosystem function. Here, I note that attributes of the predator, the prey, and the environment are all important in determining the strength of antipredator responses, and I develop hypotheses for the ways that prey functional traits might influence the magnitude of risk effects. In particular, I consider the following attributes of prey: group size and dilution of direct predation risk, the degree of foraging specialization, body mass, and the degree to which direct predation is additive vs. compensatory. Strong tests of these hypotheses will require continued development of methods to identify and quantify the fitness costs of antipredator responses in wild populations.     

The strategy of fly-and-forage migration,illustrated for the osprey (<Emphasis Type="Italic">Pandion haliaetus</Emphasis>)

Migrating birds often alternate between flight steps, when distance is covered and energy consumed, and stopover periods, when energy reserves are restored. An alternative strategy is fly-and-forage migration, useful mainly for birds that hunt or locate their prey in flight, and thus, enables birds to combine foraging with covering migration distance. The favourability of this strategy in comparison with the traditional stopover strategy depends on costs of reduced effective travel speed and benefits of offsetting energy consumption during migration flights. Evaluating these cost-benefit effects, we predict that fly-and-forage migration is favourable under many conditions (increasing total migration speed), both as a pure strategy and in combination with stopover behaviour. We used the osprey (Pandion haliaetus) as test case for investigating the importance of this strategy during spring and autumn migration at a lake in southern Sweden. The majority, 78%, of passing ospreys behaved according to the fly-and-forage migration strategy by deviating from their migratory track to visit or forage at the lake, while 12% migrated past the lake without response, and 10% made stopovers at the lake. Foraging success of passing ospreys was almost as good as for birds on stopover. Timing of foraging demonstrated that the birds adopted a genuine fly-and-forage strategy rather than intensified foraging before and after the daily travelling period. We predict that fly-and-forage migration is widely used and important among many species besides the osprey, and the exploration of its occurrence and consequences will be a challenging task in the field of optimal migration.     

Effects of predation risk and hunger on the behaviour of two species of tadpoles

Predation and hunger are threats for most organisms, and appropriate behavioural responses to both factors should be shaped by natural selection. In combination, however, the behavioural demands of predation avoidance and effective foraging often cannot be satisfied at the same time and lead to a conflict within organisms. We examined the behavioural responses of two closely-related species of tadpoles, Rana lessonae and R. esculenta, to simulated predation by fish and hunger. Tadpoles, hatched and reared in the laboratory, were tested in a three-way factorial (predation risk × hunger × species) experiment with four predation levels and four hunger levels. Both species decreased their swimming activity with increasing predation risk. Predation risk did not influence the amount of activity time invested in feeding but caused the tadpoles to spend less time in patches with food. Refuges were not used to avoid predation. R. esculenta was more sensitive to predation risk than R. lessonae. Hunger increased both the activity of tadpoles and the amount of activity time invested in feeding, thus indicating an increased energy intake. No interactions were observed between predation risk and hunger. These results show that tadpoles possess genetically-based behavioural mechanisms that allow them to respond in a graded manner to predation and hunger. However, they did not balance the two conflicting demands of predation avoidance and effective foraging; the two mechanisms appeared to act independently. Correspondence to: R.D. Semlitsch     

Predation risk and the territory value of cover: an experimental study

In an experiment on territorial brown trout (Salmo trutta), we addressed the novel hypothesis that protective cover increases the value of a territory in relation to the perceived level of predation risk. We predicted that territory holders should invest more resources defending territories with cover than territories without cover, and that defence should increase as predation risk increases. First, trout were allowed to establish ownership in territories with or without overhead cover. Second, predation risk was manipulated by simulating aerial predator attacks in half of the territories of each type, whereupon the preference for cover was estimated. Third, owners of the four types of territories were staged in dyadic contests against size-matched intruders. Territory owners showed a preference for cover, which increased further after simulated predator attacks. In subsequent contests, conflicts over territories with cover were settled faster than conflicts over territories without cover, which may suggest that the value of cover increases the motivational asymmetry between owner and intruder. Consistent with our hypothesis, owners of territories with cover were much more aggressive if they had been subjected to predator attacks the day before the conflict. These results suggest that territory owners are able to estimate the value of protective cover in response to variation in the level of predation risk in the habitat.Communicated by J. Krause     

Management of fat reserves in tufted titmice (Parus bicolor ): evidence against a trade-off with food hoards

Caching species can manage their energy supply by adjusting body fat, number of caches, or both. It has been hypothesized that because body fat has a higher fitness cost than caches, small food-hoarding birds respond to increased starvation risk by increasing the number of their caches rather than their fat load. This hypothesis predicts that when birds cannot cache they should compensate for the loss of external energy storage by (1) shifting the time of their daily body mass accumulation toward earlier in the day and (2) increasing the overall level of their fat reserves. During the winter of 1995–1996, we tested these predictions with a caching species, the tufted titmouse (Parus bicolor). Each of six experimental birds was fed a diet of uncachable sunflower seed powder for 6 days, preceded and followed by 6-day control periods during which they were fed cachable sunflower seeds. The daily pattern of body mass gain was unaffected by the opportunity to cache. Furthermore, when unable to cache, the birds did not increase either their mean daily body mass, body mass in the middle of the day, or evening body mass compared to the two control periods. These results argue against the hypothesis of a trade-off between fat reserves and food caches in tufted titmice, and suggest that fat reserves are managed independently of external food caches. Received: 9 April 1997 / Accepted after revision: 30 August 1997     

Habitat use by juvenile Atlantic cod (Gadus morhua) in the presence of an actively foraging and non-foraging predator

Experiments were conducted in the autumn and winter of 1992/93 to examine habitat use by juvenile (age 0+) Atlantic cod, Gadus morhua L., before, during and following exposure to a passive or actively foraging predator (age 3+ cod). Experiments presented groups of juvenile cod (n=5 fish/group) with one of two combinations of three substrates; (1) gravel, sand, and a patch of artificial kelp (kelp), or (2) cobble, sand, and kelp. Cobble is known to provide juvenile cod with a refuge from predation. Kelp was used to test the hypothesis that juvenile cod associate with fleshy macroalgae in nature because of the safety it provides from predators. There was little difference in habitat use by juvenile cod before, during or following exposure to a passive predator. Under these conditions, juvenile cod appeared to prefer finer grained mineral substrates and avoided the kelp. The extent of the juvenile response to a passive predator was to avoid the predator's location in the experimental tank. In contrast, juvenile cod showed a significant shift in habitat use when exposed to an actively foraging predator, hiding in cobble or, when cobble was not available, in kelp. Use of both these habitats resulted in a significant reduction in predation risk to the juvenile cod. Our results suggest that: (1) an association with kelp provides safety from predation to juvenile cod, and (2) juvenile cod are capable for assessing the risk a predator represents and adjust their response accordingly.