Influence of environment structure and food availability on the foraging behaviour of the laboratory rat

According to the optimal foraging theory, an animal is expected to enter into a given activity depending on associated costs and benefits. In line with this assumption, numerous studies have suggested that energetic reward is balanced by predation risk in foraging decisions. Therefore, the use of information about indirect cues of predation risk such as physical structure (e.g. cover, escape substrate) can give individuals a selective advantage. We studied foraging behaviour in the laboratory rat in an experimental maze; it allowed us to vary two environmental parameters: food availability and physical structure. In a first experiment, rats were offered a choice between two areas only differing in cover density. In a second experiment, the two areas only differed in food density. In a third experiment, we crossed both parameters. Our results showed that high “cover” patch was preferentially exploited (experiment 1) and that rats foraged more in the high food density patch (experiment 2). The last experiment showed that rats partially trade-off between cover density and food availability, even if the safest area was still preferred. Therefore, we suggest that foraging decisions depend primarily on safety needs, rather than food availability, at least when animals are not severely food-deprived.     

Avian daily foraging patterns: Effects of digestive constraints and variability

Summary Birds show a typical daily pattern of heavy morning and secondary afternoon feeding. We investigate the pattern of foraging by a bird that results in the lowest long-term rate of mortality. We assume the following: mortality is the sum of starvation and predation. The bird is characterized by two state variables, its energy reserves and the amount of food in its stomach. Starvation occurs during the day if the bird's reserves fall to zero. The bird starves during the night if the total energy stored in reserves and the stomach is less than a critical amount. The probability that the bird is killed by a predator is higher if the bird is foraging than if it is resting. Furthermore, the predation risk while foraging increases with the bird's mass. From these assumptions, we use dynamic programming techniques to find the daily foraging routine that minimizes mortality. The principal results are (1) Variability in food finding leads to routines with feeding concentrated early in the day, (2) digestive constraints cause feeding to be spread more evenly through the day, (3) even under fairly severe digestive constraints, the stomach is generally not full and (4) optimal fat reserve levels are higher in more variable environments and under digestive constraints. This model suggests that the characteristic daily feeding pattern of small birds is not due to digestive constraints but is greatly influenced by environmental variability.     

Evaluation of predation risk in the collectively foraging termite Macrotermes bellicosus

Summary. Few studies have investigated foraging decisions in collectively foraging social insects with no studies in termites. In termites predation is assumed to be a key mortality factor. Therefore, we experimentally investigated the role of predation pressure in foraging decisions of the fungus cultivating, mound building termite Macrotermes bellicosus in two habitats of the Comoé National Park (Ivory Coast). We used the indirect approach of measuring the Giving up Density (GUD), which is the amount of food left when individuals stop foraging in a food patch, whilst experimentally varying predation pressure. Three different conditions were examined: (a) natural predation, (b) no predation, and (c) experimental predation through artificial removal of termites. In the shrub savanna, foraging termites responded to increasing predation with increasing GUDs. By contrast, in the gallery forest, there was no gradual response. Instead termites abandoned a food patch immediately after an attack by predators. Without predation GUDs were lower in the savanna than in the gallery forest indicating that food had a higher value in the former habitat. This, together with the differential behavioral responses to predation, was in accordance with high availability of food in the gallery forest and a limited supply of food in the savanna. Thus, according to our results termites traded off predation pressure differently, according to the availability of food in both habitats.     

Behavioural Response of Juvenile Bluegill Sunfish to Variation in Predation Risk and Food Level

The behavioural response of juvenile bluegill sunfish (Lepomis macrochirus) to predation risk when selecting between patches of artificial vegetation differing in food and stem density was investigated. Bluegill foraging activity was significantly affected by all three factors. Regardless of patch stem density or risk of predation bluegills preferred patches with the highest prey number. During each trial bluegill foraging activity was clearly divided into a between- and within-patch component. In the presence of a predator bluegills reduced their between-patch foraging activity by an equivalent amount regardless of patch stem density or food level, apparently showing a risk-adjusting behavioural response to predation risk. Within patches, however, foraging activity was affected by both food level and patch stem density. When foraging in a patch offering a refuge from predation, the presence of a predator had no effect on bluegill foraging activity within this patch. However, if foraging in a patch with only limited refuge potential, bluegill foraging activity was reduced significantly in the presence of a predator. Further, this reduction was significantly greater if the patch contained a low versus a high food level, indicating a risk-balancing response to predation with respect to within-patch foraging activity. Both these responses differ from the risk-avoidance response to predation demonstrated by juvenile bluegills when selecting among habitats. Therefore, our results demonstrate the flexibility of juvenile bluegill foraging behaviour.     

Foraging Patch Selection in Winter: A Balance between Predation Risk and Thermoregulation Benefit

In winter, foraging activity is intended to optimize food search while minimizing both thermoregulation costs and predation risk. Here we quantify the relative importance of thermoregulation and predation in foraging patch selection of woodland birds wintering in a Mediterranean montane forest. Specifically, we account for thermoregulation benefits related to temperature, and predation risk associated with both illumination of the feeding patch and distance to the nearest refuge provided by vegetation. We measured the amount of time that 38 marked individual birds belonging to five small passerine species spent foraging at artificial feeders. Feeders were located in forest patches that vary in distance to protective cover and exposure to sun radiation; temperature and illumination were registered locally by data loggers. Our results support the influence of both thermoregulation benefits and predation costs on feeding patch choice. The influence of distance to refuge (negative relationship) was nearly three times higher than that of temperature (positive relationship) in determining total foraging time spent at a patch. Light intensity had a negligible and no significant effect. This pattern was generalizable among species and individuals within species, and highlights the preponderance of latent predation risk over thermoregulation benefits on foraging decisions of birds wintering in temperate Mediterranean forests.     

Variation of within-day foraging costs in the striped mouse (Rhabdomys pumilio)

Foraging behavior is influenced by spatial and temporal habitat heterogeniety. Here we report on within-day foraging and perceived risk of predation by the striped mouse (Rhabdomys pumilio) in a grassland savannah with wooded “islands” using giving-up densities (GUD, amount of food left behind in depletable food patches). Higher GUDs correspond to higher forging costs. GUDs were measured six times per day at 2-h intervals from paired stations along fern–grass habitat boundaries at 3 and 6 m distances from 10 wooded islands. R. pumilio's GUDs varied significantly over the course of the day with highest GUDs during the afternoon hours of 1–3 pm, and lowest between 7 and 9 am in the morning. The same pattern was consistent for both habitats (fern and grass) and distances from the wooded islands. GUDs decreased with distance from the woody islands in both fern and grass habitats and were significantly lower in the fern habitat. This activity pattern suggests that R. pumilio responds to a spectrum of spatially and temporally varying risks from a variety of predators including aerial predators that increase risk as they make use of mid-day thermals.     

Why do hoarding birds gain fat in winter in the wrong way? Suggestions from a dynamic model

In winter, small birds should be fat to avoid starvation andlean and agile to escape predators. This means that they facea trade-off between the costs and benefits of carrying fat reserves.Every day they must gain enough fat to survive the coming night.Food-hoarding species can afford to carry less fat than nonhoardersbecause they can store energy outside the body. Furthermore, hoardersshould avoid carrying excessive fat during the day because theycan gain fat fast by retrieving food late in the afternoon.With no stored supplies, nonhoarders face more unpredictableaccess to food, and they should start gaining fat earlier inthe day. The predicted pattern is then that nonhoarders gainfat early and that hoarders gain fat late in the day. Recent fielddata show the opposite pattern: hoarders gain relatively morefat reserves in the morning than nonhoarders do. Using a dynamicmodel that mimics the conditions in a boreal winter forest,I investigated under which conditions this pattern will arise.The only assumption of those investigated that produced thispattern was to relax the effect of mass-dependent predation risk.I did this by introducing a limit under which fat reserves didnot affect predation risk. Hoarders then started the day bygaining fat in the morning. Later, when they had reached a safer(but still not risky) level, they switched to hoarding. Thepattern I searched would only occur if either not all food waspossible to store, or if retrieval gave less energy than foragingin good weather conditions. If I assumed that low levels ofbody fat also increased predation risk, hoarders would cachein the morning when they carried least fat. I discuss empiricalevidence for how body fat affects predation risk. In summary,the factors that produced the pattern I searched were a changein the predation-mortality function combined with restrictions onhoarding.     

Meals for two: foraging activity of the butterflyfish Chaetodon striatus (Perciformes) in southeast Brazil.

The banded butterflyfish (Chaetodon striatus) from the tropical and subtropical western Atlantic is a territorial, diurnal forager on benthic invertebrates. It is usually seen moving singly or in pairs, a few meters above the sea floor. We studied the foraging activity of C. striatus on rocky reefs in southeastern Brazil. This fish spent about 11 h and 30 min per day on feeding activities, and preferred colonies of non-scleratinian anthozoans over sandy and rocky substrata while foraging. The lowest feeding rates were recorded in the early morning and late afternoon, but we found no further differences between feeding rates throughout the day. We also found no differences between the feeding rates of paired and single individuals.     

Foraging patterns of voles at heterogeneous avian and uniform mustelid predation risk

Temporal variation of antipredatory behavior and a uniform distribution of predation risk over refuges and foraging sites may create foraging patterns different from those anticipated from risk in heterogenous habitats. We studied the temporal variation in foraging behavior of voles exposed to uniform mustelid predation risk and heterogeneous avian predation risk of different levels induced by vegetation types in eight outdoor enclosures (0.25 ha). We manipulated mustelid predation risk with weasel presence or absence and avian predation risk by reducing or providing local cover at experimental food patches. Foraging at food patches was monitored by collecting giving-up densities at artificial food patches, overall activity was automatically monitored, and mortality of voles was monitored by live-trapping and radiotracking. Voles depleted the food to lower levels in the sheltered patches than in the exposed ones. In enclosures with higher avian predation risk caused by lower vegetation height, trays were depleted to lower levels. Unexpectedly, voles foraged in more trays and depleted trays to lower levels in the presence of weasels than in the absence. Weasels match their prey's body size and locomotive abilities and therefore increase predation risk uniformly over both foraging sites and refuge sites that can both be entered by the predator. This reduces the costs of missing opportunities other than foraging. Voles changed their foraging strategy accordingly by specializing on the experimental food patches with predictable returns and probably reduced their foraging in the matrix of natural food source with unpredictable returns and high risk to encounter the weasel. Moreover, after 1 day of weasel presence, voles shifted their main foraging activities to avoid the diurnal weasel. This behavior facilitated bird predation, probably by nocturnal owls, and more voles were killed by birds than by weasels. Food patch use of voles in weasel enclosures increased with time. Voles had to balance the previously missed feeding opportunities by progressively concentrating on artificial food patches.     

Personality predicts individual responsiveness to the risks of starvation and predation

Theory suggests that individual personality is tightly linked to individual life histories and to environmental variation. The reactive-proactive axis, for example, is thought to reflect whether individuals prioritize productivity or survival, mutually exclusive options that can be caused by conflicts between foraging and anti-predation behaviour. Evidence for this trade-off hypothesis, however, is limited. Here, we tested experimentally whether exploration behaviour (EB), an assay of proactivity, could explain how great tits (Parus major) respond to changes in starvation and predation risk. Individuals were presented with two feeders, holding good or poor quality food, which interchanged between safe and dangerous positions 10 m apart, across two 24 h treatments. Starvation risk was assumed to be highest in the morning and lowest in the afternoon. The proportion of time spent feeding on good quality food (PTG) rather than poor quality food was repeatable within treatments, but individuals varied in how PTG changed with respect to predation- and starvation-risk across treatments. This individual plasticity variation in foraging behaviour was linked to EB, as predicted by the reactive-proactive axis, but only among individuals in dominant social classes. Our results support the trade-off hypothesis at the level of individuals in a wild population, and suggest that fine-scale temporal and spatial variation may play important roles in the evolution of personality.     

Time–Place Learning Based on Food but not on Predation Risk in a Fish, the Inanga (Galaxias maculatus)

Time–place learning, or the ability to learn to be in different places at different times of day, is already known to occur in response to daily spatio-temporal patterns of food availability. However, the ability to learn daily patterns of predation risk and move between areas at the right time of day in order to avoid predation has never been tested. This study asked whether inangas, Galaxias maculatus , are capable of time–place learning based on food availability only, predation risk only, or the antagonistic combination of food availability and predation risk. Shoals of five inangas were kept in aquaria partially divided into a right and left section. Every day they were exposed to a stimulus on one side in the morning and on the other side in the afternoon. Depending on the experiment, the stimulus could be two deliveries of food, two simulated heron strikes, or both of the above within the same hour. After 14 d the stimuli were not given and the position of the fish was noted in both the morning and the afternoon. The majority of the fish learned to switch sides at the correct daily time in order to get food, but they remained on the same side at both daily times in response to either predation risk alone or the combination of predation risk and food. It seems that the potential for time–place learning based on predation risk is less than that based on food, and that predation risk can even curtail the expression of time–place learning based on food. Fish may resort to other tactics, such as shoaling and reduced movement, in response to predation risk. Daily habitat shifts could still be present in nature and rooted in the avoidance of predation, but instead of being the direct result of learning they would be mostly innate.     

Fat reserves and perceived predation risk in the great tit, Parus major

The fat reserves of small birds are built up daily as insurance against starvation. They are believed to reflect a trade-off between the risks of starvation and predation such that in situations of high predation risk birds are expected either to reduce their fat reserves in response to mass-dependent predation risk or to increase them in response to foraging interruptions. We assessed the effect on fat reserves of experimentally altering the perceived (but not the actual) risk of predation of wild great tits at a winter feeding site. The perceived predation risk was alternated between 'safe' and 'risky'. Increasing the perceived risk of predation involved 'swooping' a model sparrowhawk over the feeder at four unpredictable times each day using a remote mechanism We produce evidence that the experiment was suceessfull in altering the perceived risk of predation. As predicted from the hypothesis of mass-dependent predation risk, great tits (Parus major) carried significantly reduced fat reserves during the 'risky' treatment. Furthermore, dominant individuals were able to reduce their reserves more than subordinates. As birds returned to feeders within seconds after a predator 'attack', the reduction in fat reserves cannot be attributed to an interruption in feeding.     

Titrating the cost of plant toxins against predators: determining the tipping point for foraging herbivores

1. Foraging herbivores must deal with plant characteristics that inhibit feeding and they must avoid being eaten. Principally, toxins limit food intake, while predation risk alters how long animals are prepared to harvest resources. Each of these factors strongly affects how herbivores use food patches, and both constraints can pose immediate proximate costs and long-term consequences to fitness. 2. Using a generalist mammalian herbivore, the common brushtail possum (Trichosurus vulpecula), our aim was to quantitatively compare the influence of plant toxin and predation risk on foraging decisions. 3. We performed a titration experiment by offering animals a choice between non-toxic food at a risky patch paired with food with one of five toxin concentrations at a safe patch. This allowed us to identify the tipping point, where the cost of toxin in the safe food patch was equivalent to the perceived predation risk in the alternative patch. 4. At low toxin concentration, animals ate more from the safe than the risky patch. As toxin concentration increased at the safe patch, intake shifted until animals ate mainly from the risky patch. This shift was associated with behavioural changes: animals spent more time and fed longer at the risky patch, while vigilance increased at both risky and safe patches. 5. Our results demonstrate that the variation in toxin concentration, which occurs intraspecifically among plants, can critically influence the relative cost of predation risk on foraging. We show that herbivores quantify, compare and balance these two different but proximate costs, altering their foraging patterns in the process. This has potential ecological and evolutionary implications for the production of plant defence compounds in relation to spatial variation in predation risk to herbivores.     

捕食风险对不同饥饿状态下艾虎取食行为的影响

在室内条件下,将大鵟作为艾虎的天敌动物,通过双通道选择实验确定6 只成体艾虎在3 个捕食风险水平和4 种饥饿状态条件下的取食行为,探讨艾虎在取食过程中对饥饿风险与捕食风险的权衡策略。研究结果表明:在无捕食风险存在时,艾虎被剥夺食物0 d 和1 d 后对食物量不同的两个斑块中的取食量和利用频次均无明显不同(P > 0. 05),但对高食物量斑块的利用时间均明显高于低食物量斑块的(P <0.05),而艾虎被剥夺食物2 d和3 d后对高食物量斑块中的取食量和利用时间均明显高于低食物量斑块中的(P < 0.05),但在利用频次上均无明显差异(P > 0.05)。在面临低风险时,艾虎在4 种饥饿状态下均只利用无天敌动物存在的低食物量斑块,而基本不利用有天敌动物存在的高食物量斑块。在面临高风险时,艾虎不得不利用有天敌动物存在的食物斑块,被剥夺食物0 d 时艾虎对无风险、无食物量斑块的利用时间基本相同于对高风险、有食物量斑块的利用时间(P>0.05),而被剥夺食物1d、2 d 和3 d 后艾虎对高风险、有食物量斑块的利用时间明显高于无风险、无食物量斑块的(P< 0. 05)。在相同风险条件下,随着饥饿程度增加,艾虎在斑块中的取食量均明显增加(P< 0.05),而对斑块的利用时间和利用频次明显降低(P<0.05)。在相同的饥饿状态下,不同风险水平时,艾虎在斑块中的取食量无明显的差异(P>0.05),但在低风险和高风险时对斑块的利用时间和频次均明显低于无风险时的(P <0.05)。以上结果说明艾虎能够根据食物摄取率和自身的能量需求在捕食风险和饥饿风险之间做出权衡,当饥饿风险小于捕食风险时,艾虎趋于躲避捕食风险,当饥饿风险大于捕食风险时,艾虎趋于面对捕食风险,所采用的取食策略是减少活动时间和能量消耗,最大程度地提高单位时间内获得的能量。     

Daily patterns of body mass gain in four species of small wintering birds

Theoretically, the trajectories describing the daily accumulation of body reserves are expected to differ between bird species in relation to whether or not they hoard food. To carry reserves on the body may be costly and hoarding species can be expected to hoard food early in the day when light and retrieve it in the afternoon, with a concomitant rapid increase in body mass. Also, the increased food predictability resulting from being able to consume hoarded food late in the day should lead to a relatively faster gain in body reserves in the afternoon in hoarding species compared to non-hoarders. Non-hoarders may have to hedge against possible afternoon losses of foraging opportunities by accumulating more reserves early in the day.
In this study the daily patterns of body mass gain in four small bird species resident during winter in Scandinavia are described. Individually known birds were trained to come to a permanent feeder and their body masses were recorded every hour throughout the day with a remote-controlled balance. The hoarding willow tit Parus montanus , marsh tit P. palustris and European nuthatch Sitta europaea all displayed the most rapid gain in body mass in the early hours of the day. After the initial burst in the morning, reserves were accumulated at a roughly constant rate for the remainder of the day. In contrast, the non-hoarding great tit P. major apparently gained body reserves at a more even rate. The daily pattern of body mass gain found in the hoarding species differs from prevailing theoretical predictions, whereas the pattern in the non-hoarding great tit is in a better agreement with theory, from which this pattern has been predicted repeatedly.     

Ectoparasites: are they the proximate cause of cleaning interactions?

We tested the importance of ectoparasites in cleaning symbioses by comparing the activity of Caribbean cleaning gobies ( Elacatinus evelynae ) and of their clients during three daily periods (early morning, midday and late afternoon) in which ectoparasite availability varied naturally. Emergence from the benthos of gnathiid isopod larvae, the main target of cleaning goby predation, was higher at night, when cleaners are inactive, than during the day. Overall ectoparasite loads also tended to be higher on clients in the morning. This coincided with higher rates of visits to cleaning stations by client fish in the morning than at midday, but high rates of client visits were also recorded in the late afternoon. Clients were more likely to adopt stereotypical incitation poses, which increase the likelihood of being cleaned, in the morning than later in the day. Inspection bouts by cleaning gobies were longest in the morning. Cleaner and client behaviours therefore change predictably in response to natural diurnal variation in ectoparasite availability. These results add to a growing number of studies supporting the idea that cleaning symbioses are mutualisms dependent on ectoparasite removal.     

The dilemma of foraging herbivores: dealing with food and fear

For foraging herbivores, both food quality and predation risk vary across the landscape. Animals should avoid low-quality food patches in favour of high-quality ones, and seek safe patches while avoiding risky ones. Herbivores often face the foraging dilemma, however, of choosing between high-quality food in risky places or low-quality food in safe places. Here, we explore how and why the interaction between food quality and predation risk affects foraging decisions of mammalian herbivores, focusing on browsers confronting plant toxins in a landscape of fear. We draw together themes of plant–herbivore and predator–prey interactions, and the roles of animal ecophysiology, behaviour and personality. The response of herbivores to the dual costs of food and fear depends on the interplay of physiology and behaviour. We discuss detoxification physiology in dealing with plant toxins, and stress physiology associated with perceived predation risk. We argue that behaviour is the interface enabling herbivores to stay or quit food patches in response to their physiological tolerance to these risks. We hypothesise that generalist and specialist herbivores perceive the relative costs of plant defence and predation risk differently and intra-specifically, individuals with different personalities and physiologies should do so too, creating individualised landscapes of food and fear. We explore the ecological significance and emergent impacts of these individual-based foraging outcomes on populations and communities, and offer predictions that can be clearly tested. In doing so, we provide an integrated platform advancing herbivore foraging theory with food quality and predation risk at its core.     

Ant assemblage composition explains high predation pressure on artificial caterpillars during early night

1. Predator–prey interactions, especially those involving herbivorous insects, are of great importance in maintaining biodiversity. Predation pressure varies temporally in response to prey availability and activity. However, little is known about the patterns and drivers of fluctuations in predation pressure at fine temporal scales. 2. Artificial caterpillars (placed on plant leaves at breast height) were used to assess changes in predation pressure across four time intervals of the day in a monsoonal tropical rainforest in south-west China. The study examined how assemblage composition of arboreal ants, the dominant predators, changed across the same time intervals. The potential linkages between biotic (arboreal ants) and abiotic (temperature and light intensity) factors with predation rate were evaluated. 3. Predation rate on caterpillars during the early part of the night (19.00–01.00 hours) was significantly higher than in the morning, afternoon, or late night. Ant assemblage composition, rather than species richness or total abundance, best explained the variations in predation rate on artificial caterpillars. 4. The results help to strengthen understanding of trophic interactions by demonstrating that predation pressure fluctuates at finer timescales than previously tested, and that a particular set of ant species may play major roles in predation on caterpillars and possibly other organisms.     

Foraging and habitat use of common duikers,Sylvicapra grimmia,in a heterogeneous environment within the Soutpansberg,South Africa

We investigated aspects of the foraging behaviour and activity patterns of free‐ranging common duikers (Sylvicapra grimmia) within the Soutpansberg, South Africa. We used giving‐up densities (GUD) and camera traps to test for habitat selection and patch‐use behaviour by common duikers inhabiting a grassland containing distinct ‘islands' of woody vegetation. Foraging in or around a wooded island was affected by its surrounding vegetation. GUDs were significantly lower in portions dominated by tall grass and scattered ferns and highest in areas with open short grass and thick fern. Using grids of 5 × 5 stations, we mapped the duikers' foraging on a larger scale that incorporated neighbouring rocky hillsides. The duikers preferred feeding in areas with tall grass and scattered fern (sufficient cover and escape routes), followed by the wooded islands and thick fern (lack of sightlines/escape routes and presence of predator‐ambush sites), whereas little foraging occurred at the edges and rocky areas (hard substrate that impede escape potential). Photos (total 873) revealed solitary activity, highest in the late afternoon. Photos of vigilant individuals were mostly from rocky and fern habitats. Our results suggest that the duikers allocated their feeding efforts, activity and vigilance patterns to attune to their perceived risk of predation within their heterogeneous environment.     

Patch use in time and space for a meso-predator in a risky world

Predator–prey studies often assume a three trophic level system where predators forage free from any risk of predation. Since meso-predators themselves are also prospective prey, they too need to trade-off between food and safety. We applied foraging theory to study patch use and habitat selection by a meso-predator, the red fox. We present evidence that foxes use a quitting harvest rate rule when deciding whether or not to abandon a foraging patch, and experience diminishing returns when foraging from a depletable food patch. Furthermore, our data suggest that patch use decisions of red foxes are influenced not just by the availability of food, but also by their perceived risk of predation. Fox behavior was affected by moonlight, with foxes depleting food resources more thoroughly (lower giving-up density) on darker nights compared to moonlit nights. Foxes reduced risk from hyenas by being more active where and when hyena activity was low. While hyenas were least active during moon, and most active during full moon nights, the reverse was true for foxes. Foxes showed twice as much activity during new moon compared to full moon nights, suggesting different costs of predation. Interestingly, resources in patches with cues of another predator (scat of wolf) were depleted to significantly lower levels compared to patches without. Our results emphasize the need for considering risk of predation for intermediate predators, and also shows how patch use theory and experimental food patches can be used for a predator. Taken together, these results may help us better understand trophic interactions.