Sampling behavior of starlings foraging in simple patchy environments

Summary Starlings were allowed to forage in patchy laboratory environments where patches contained either zero or a fixed number of prey. The condition of a given patch (prey or no prey) could only be determined from information gained while exploiting the patch. A starling's task was to determine to what extent to sample an apparently prey-less patch before giving it up as such, in a way which maximizes long-term energy intake rate. The simple model presented to predict the optimal sampling solutions was qualitatively but not quantitatively supported by the data. The main discrepancy was in the fact that an apparently prey-less patch should have been sampled to a fixed extent before leaving, whereas a distribution of sampling behavior was actually observed. The qualitative agreement was very good, however, as the modes of the observed sampling distributions often corresponded to the predicted optimal sampling solutions. Starlings seem to possess a patch-sampling ability which, at least for those simple situations analyzed, can lead to an efficient foraging strategy.     

Optimal movement strategies for social foragers in unpredictable environments

Spatial movement models often base movement decision rules on traditional optimal foraging theories, including ideal free distribution (IFD) theory, more recently generalized as density-dependent habitat selection (DDHS) theory, and the marginal value theorem (MVT). Thus optimal patch departure times are predicted on the basis of the density-dependent resource level in the patch. Recently, alternatives to density as a habitat selection criterion, such as individual knowledge of the resource distribution, conspecific attraction, and site fidelity, have been recognized as important influences on movement behavior in environments with an uncertain resource distribution. For foraging processes incorporating these influences, it is not clear whether simple optimal foraging theories provide a reasonable approximation to animal behavior or whether they may be misleading. This study compares patch departure strategies predicted by DDHS theory and the MVT with evolutionarily optimal patch departure strategies for a wide range of foraging scenarios. The level of accuracy with which individuals can navigate toward local food sources is varied, and individual tendency for conspecific attraction or repulsion is optimized over a continuous spectrum. We find that DDHS theory and the MVT accurately predict the evolutionarily optimal patch departure strategy for foragers with high navigational accuracy for a wide range of resource distributions. As navigational accuracy is reduced, the patch departure strategy cannot be accurately predicted by these theories for environments with a heterogeneous resource distribution. In these situations, social forces improve foraging success and have a strong influence on optimal patch departure strategies, causing individuals to stay longer in patches than the optimal foraging theories predict.     

Searching behavior and use of sampling information by free-ranging bison (Bos bison)

I examined the searching behavior of free-ranging plains bison (Bos bison bison) in their natural habitat, and determined whether their assessment of food patch quality was influenced by the short-term sampling information acquired during search. Bison used area-concentrated search during their winter foraging activity. Their movements between areas of suitable food patches were influenced by local environmental conditions, being sometimes less sinuous, and at other times more sinuous, than expected from a correlated random walk model. Bison also systematically avoided digging in areas where plants of low profitability lay under the snow. Where they dug, there was evidence that a bison's perception of food quality varied during a foraging bout, and was therefore influenced by short-term sampling information. After controlling for forage quality, I found that small feeding craters were more likely to be preceded by samples of high quality food patches. My observations suggest that bison take advantage of the structural characteristics of their environment during searching activity, and base foraging decisions on local rather than global availability.     

Effects of within- and among-patch experiences on the patch-leaving decision rules in an insect parasitoid

The present study aimed to address how an insect parasitoid makes patch-departure decisions from various types of host patches and how previous patch experiences in the environment modify this decision-making process. Experiments were done with the parasitic wasp Aphidius rhopalosiphi attacking the grain aphid Sitobion avenae. In the experiments, wasps were observed in a laboratory environment containing several patches of various host densities, and behavioural records were analysed using a Coxs proportional hazards model. Consideration of the effect of the within-patch experience gave a classic pattern of patch-leaving decision rules in parasitoids: A. rhopalosiphi used local information on host quality (i.e. numbers of ovipositions or rejections) and availability (i.e. patch density) to determine departure decision. However, consideration of previous patch experiences provided evidence that these departure rules are fundamentally dynamic, responding to the physiological state of the animal and the information it has about its environment. Results showed that A. rhopalosiphi decreased its tendency to leave the visited patch after an oviposition. However, when a female has already laid several other eggs in the environment, such an incremental mechanism gradually switched to a decremental one. Hence, A. rhopalosiphi responded to egg-load depletion by leaving the visited patches sooner and by depositing a smaller number of eggs in those patches, which probably led to a decreased level of superparasitism. Results also indicated that previous experiences enabled wasps to estimate spatial host distribution and then to adjust their behavioural decisions accordingly. Thus, A. rhopalosiphi was shown to adjust its patch residence time according to the quality and the number of the patches previously visited. These proximate mechanistic rules adopted by A. rhopalosiphi females are discussed in the context of general predictions from optimality models.Communicated by D. Gwynne     

Subhabitat selection by foraging threespine stickleback (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>): previous experience and social conformity

Any mechanism that allows animals to increase their foraging efficiency is likely to be selected for, including the ability to learn to recognise and subsequently discriminate between habitat types based on their profitability. In a series of laboratory studies, we manipulated prey densities across two different experimental subhabitats and demonstrated that threespine stickleback (Gasterosteus aculeatus) can develop foraging preferences for subhabitats that have previously yielded prey. Fish were not recalling the spatial location of prey patches; rather, they were discriminating between subhabitats based on foraging experience there and allocating foraging effort accordingly. Foraging preferences took around 14 days to develop, and once established, they persisted independently of experimental prey density, suggesting that fish were using experience rather than real-time sampling to select foraging grounds. When we presented focal fish with social information cues, we found that they preferentially used local enhancement and current public information cues when they conflicted with previous experience, but that they did not use prior public information. This suggests that in the presence of conspecifics, individuals prioritise social conformity over the use of private information. We discuss our results in the context of optimal foraging and the trade-offs associated with balancing conflicting private and social information.     

Travel time affects optimal diets in depleting patches

Models of prey choice in depleting patches predict an expanding specialist strategy: Animals should start as specialists on the most profitable prey and then at some point during patch exploitation switch to a generalist foraging strategy. When patch residence time is long, the switch to a generalist diet is predicted to occur earlier than when patch residence time is short. We tested these predictions under laboratory conditions using female parasitoids (Aphidius colemani) exploiting patches of mixed instars aphid hosts (Myzus persicae, L1 and L4). The duration of patch exploitation was manipulated by changing travel time between patches. As predicted, patch residence times increase with travel time between patches. Our results provide empirical support for the expanding specialist prediction: Parasitoid females specialized initially on the more profitable hosts (L4), and as the patch depleted, they switched to a generalist diet by accepting more frequently the less profitable hosts (L1). The point at which they switched from specialist to generalist occurred later when travel times and hence patch residence times were short. By affecting the patch exploitation strategy, travel time also determines the composition of hosts left behind, the “giving up composition.” The change in the relative density of remaining host types alters aphid populations’ age structure.     

A field investigation of scrounging in semipalmated sandpipers

Animals that forage in groups can produce their own food patches or scrounge the food discoveries of their companions. Mean tactic payoffs are expected to be the same at equilibrium for phenotypically equal foragers. Scrounging is also typically viewed as a risk-averse foraging strategy that provides a more even food intake rate over time. The occurrence of scrounging and the payoffs from different foraging modes have rarely been investigated in the field. Over two field seasons, I examined patch sharing in semipalmated sandpipers (Calidris pusilla) foraging on minute food items at the surface of the substrate. Birds could find patches on their own, a producing event, or join the food patches discovered by others, a scrounging event. I found that the average search time per patch did not differ between producing and scrounging but that the average time spent exploiting a patch was reduced nearly by half when scrounging. As a result, the proportion of time spent exploiting a patch, a measure of foraging payoffs, was significantly lower when scrounging. The variance in payoffs was similar for producing and scrounging. When producing their own patches, individuals that scrounged spent the same proportion of time exploiting a patch as those that only produced. However, within the same individuals, the search time for a scrounged patch was longer than the search time for a produced patch. The results show unequal payoffs for producing and scrounging in this system and suggest that low success in finding patches elicited scrounging.     

Dance precision of <Emphasis Type="Italic">Apis florea</Emphasis>—clues to the evolution of the honeybee dance language?

All honeybee species make use of the waggle dance to communicate the direction and distance to both food sources and potential new nest sites. When foraging, all species face an identical problem: conveying information about profitable floral patches. However, profound differences in nesting biology (some nest in cavities while others nest in the open, often on a branch or a cliff face) may mean that species have different requirements when dancing to advertise new nest sites. In cavity nesting species, nest sites are a precise location in the landscape: usually a small opening leading to a cavity in a hollow tree. Dances for cavities therefore need to be as precise as possible. In contrast, when the potential nest site comprises a tree or perhaps seven a patch of trees, precision is less necessary. Similarly, when a food patch is advertised, dances need not be very precise, as floral patches are often large, unless they are so far away that recruits need more precise information to be able to locate them. In this paper, we study the dance precision of the open-nesting red dwarf bee Apis florea. By comparing the precision of dances for food sources and nest sites, we show that A. florea workers dance with the same imprecision irrespective of context. This is in sharp contrast with the cavity-nesting Apis mellifera that increases the precision of its dance when advertising a potential new home. We suggest that our results are in accordance with the hypothesis that the honeybees’ dance communication initially evolved to convey information about new nest sites and was only later adapted for the context of foraging.     

Locating food in a spatially heterogeneous environment: Implications for fitness of the macrodecomposer Armadillidium vulgare (Isopoda: Oniscidea)

To assess the fitness consequences of foraging on patchy resources, consumption rates, growth rates and survivorship of Armadillidium vulgare were monitored while feeding in arenas in which the spatial distribution of patches of high quality food (powdered dicotyledonous leaf litter) was varied within a matrix of low quality food (powdered grass leaf litter). Predictions from behavioural experiments that these fitness correlates would be lower when high quality food is more heterogeneously distributed in space were tested but not supported either by laboratory or field experiments. To investigate whether A. vulgare can develop the ability to relocate high quality food patches, changes in foraging behaviour, over a comparable time period to that used in the fitness experiments, were monitored in arenas in which there was a high quality food patch in a low quality matrix. A. vulgare increased its ability to relocate the position of high quality food over time. It reduced time spent in low quality food matrices and increased time spent in high quality food patches with time after the start of the experiment. When the position of a high quality food patch was moved, the time spent in the low quality food matrix increased and less time was spent in high quality food patches, compared to arenas in which the food was not moved. The fitness benefits for saprophages of developing the ability to relocate high quality patches while foraging in spatially heterogeneous environments are discussed.     

Individual differences in sampling behaviour predict social information use in zebra finches

When animals have to decide where to forage, what to eat or with whom to mate, they can base their decisions on either socially or personally acquired information. In accordance with theoretical predictions, there is experimental evidence that animals adjust the weight they give to both sources of information depending on circumstances. Notably, several studies have demonstrated that individuals rely more on social information when personal information is difficult to acquire or unreliable, because these conditions leave them uncertain as how to behave. Yet, even when individuals are exposed to the same conditions, they generally differ widely in the value they attribute to social and personal information. These differences suggest that the tendency to rely on social information would also depend on intrinsic characteristics that affect, for instance, individual efficiency in collecting personal information. To address this issue, we conducted laboratory experiments using female zebra finches (Taeniopygia guttata) and we tested them under three consecutive conditions. First, we evaluated their reliance on social information in a mating context and in a foraging context. Then, we measured their efficiency in acquiring personal information by recording their sampling behaviour when searching for hidden food. We found that females that sampled their environment less actively consistently relied on social information to a greater extent compared with those that invested more in sampling. Contrary to what is generally assumed, then, our study demonstrates that social information use is not entirely flexible and context dependent.     

Integrated modelling of foraging behaviour, energy budget and memory properties

A predator's foraging performance is related to its ability to acquire sufficient information on environmental profitability. This process can be affected by the patchy distribution and clustering of food resources and by the food intake process dynamics.We simulated body mass growth and behaviour in a forager acting in a patchy environment with patchy distribution of both prey abundance and body mass by an individual-based model. In our model, food intake was a discrete and stochastic process and leaving decision was based on the estimate of net energy gain and searching time during their foraging activities. The study aimed to investigate the effects of learning processes and food resource exploitation on body mass and survival of foragers under different scenarios of intra-patch resource distribution.The simulation output showed that different sources of resource variability between patches affected foraging efficiency differently. When prey abundance varied across patches, the predator stayed longer in poorest patches to obtain the information needed and its performance was affected by the cost of sampling and the resulting assessment of the environment proved unreliable. On the other hand, when prey body mass, but not abundance, varied among the patches the predator was quickly able to assess local profitability. Both body mass and survival of the predator were greatly affected by learning processes and patterns of food resource distribution.     

Patch departure rules in Bumblebees: evidence of a decremental motivational mechanism

The patch living rules of a pollinator, the bumblebee Bombus terrestris L., are studied here in the framework of motivational models widely used for parasitoids: The rewarding events found during the foraging process are supposed to increase or decrease suddenly the tendency of the insect to stay in the current patch and therefore to adjust the patch residence time to the patch profitability. The foraging behaviour of these pollinators was observed in two environment types to determine their patch-leaving decisions. The rich environment was composed of male-fertile flowers, offering pollen and nectar, and the poor one of male-sterile flowers, offering little nectar and no pollen. The experimental design consisted of a patch system in which inflorescences were evenly arranged in two rows (1 m distance). Residence times of foragers inside inflorescences and rows were analysed by a Cox proportional hazards model, taking into account recent and past experience acquired during the foraging bout. Most of the results showed a decremental motivational mechanism, that is, a reduction in the residence time on the inflorescence or in the row related to exploitation of flowers within inflorescences and inflorescences within rows These results indicate that bumblebees tend to leave the patch using departure rules similar to those found in parasitoids. The results also provide information on the memory, learning and evaluating capabilities of bumblebees especially when rich and poor environments were compared. The patch-leaving mechanism suggested by this study is consistent with the central place foraging theory.     

Trunk trails and the searching strategy of a leaf-cutter ant,Atta colombica

Summary Atta colombica uses chemical mass recruitment that allows the rapid exploitation of resources. Most foragers thus search only within patches. Accumulation of extra foragers at patches results in sampling of alternate food items and area-restricted search as patch resources are depleted.Individual workers have a higher probability of removing a leaf fragment the earlier they arrive at a bait. Workers that arrive when much of the resource is gone travel further on the bait (within the patch) but do not spend significantly more time at the patch. They give up after 50–80s.Foraging effort is centered on the extensive trail system, not on the nest a predicted by time and energy foraging models. Search effort is also trail centered. The probability that an item will be discovered decreases with distance from the trail and increasing litter depth. Trail traffic and trail quality together mave no significant effect although this may be because they act antagonistically.Economic considerations predict that trials should be built to high quality and very productive sites. If trails are built as a result of recruitment and recruitment reflects patch quality and productivity, characteristics of forage sites are physically embodied in the trail system.Leaf cutter foraging is better understood as a long term optimization that effectively exploits resources over the lifetime of the colony than as prudent predation that husbands resources.     

Lévy flight patterns are predicted to be an emergent property of a bumblebees’ foraging strategy

Bumblebees forage uninterrupted for long periods of time because they are not distracted by sex or territorial defense and have few predators. This has led to a long running debate about whether bumblebees forage optimally. This debate has been enriched by the possibility that bumblebees foraging within clover patches have flight patterns that can be approximated by Lévy flights. Such flight patterns optimise the success of random searches. Bumblebees foraging within a flower patch tend to approach the nearest flower but then often depart without landing or probing it if it has been visited previously; unvisited flowers are not rejected in this manner. Here, this foraging behaviour has been replicated in numerical simulations. Lévy flight patterns are found to be an inconsequential emergent property of a bumblebees’ foraging behaviour. Lévy flights are predicted to emerge when bees reject at least 99% of previously visited flowers. A foraging bumblebee can certainly empty a clover flower head of nectar in one visit, but lower rates of rejection are observed for many other flowers. These findings suggest that Lévy flight patterns in foraging bumblebees are rare and specific to a few flower species and that if they exist, then they are not part of an innate, evolved optimal searching strategy.     

Foraging bumblebees do not rate social information above personal experience

Foraging animals can acquire new information about food sources either individually or socially, but they can also opt to rely on information that they have already acquired, termed “personal information”. Although social information can provide an adaptive shortcut to new resources, recent theory predicts that investing too much time in acquiring new information can be detrimental. Here, we investigate whether foraging bumblebees (Bombus terrestris) strategically prioritize personal information unless there is evidence of environmental change. All bees in our study had personal information that one species of artificial flower was rewarding, and bees in the scent group then experienced social information about an alternative-scented species inside the nest, while a control group did not. On their next foraging bout, bees in both groups overwhelmingly used personal information when deciding where to forage. When bees subsequently learnt that the rewards offered by their preferred species had dwindled, bees that had social information were no quicker to abandon their personal information than control bees, but once they had sampled the alternative flowers, they showed greater commitment to that species than control bees. Thus, we found no evidence that social information is particularly important when personal information fails to produce rewards (a “copy when established behaviour is unproductive” strategy). Instead, bees used social information specifically to complement personal information.     

Look before you leap: is risk of injury a foraging cost?

Theory states that an optimal forager should exploit a patch so long as its harvest rate of resources from the patch exceeds its energetic, predation, and missed opportunity costs for foraging. However, for many foragers, predation is not the only source of danger they face while foraging. Foragers also face the risk of injuring themselves. To test whether risk of injury gives rise to a foraging cost, we offered red foxes pairs of depletable resource patches in which they experienced diminishing returns. The resource patches were identical in all respects, save for the risk of injury. In response, the foxes exploited the safe patches more intensively. They foraged for a longer time and also removed more food (i.e., had lower giving up densities) in the safe patches compared to the risky patches. Although they never sustained injury, video footage revealed that the foxes used greater care while foraging from the risky patches and removed food at a slower rate. Furthermore, an increase in their hunger state led foxes to allocate more time to foraging from the risky patches, thereby exposing themselves to higher risks. Our results suggest that foxes treat risk of injury as a foraging cost and use time allocation and daring—the willingness to risk injury—as tools for managing their risk of injury while foraging. This is the first study, to our knowledge, which explicitly tests and shows that risk of injury is indeed a foraging cost. While nearly all foragers may face an injury cost of foraging, we suggest that this cost will be largest and most important for predators.     

Predation risk and foraging behavior of the hoary marmot in Alaska

Summary I observed hoary marmots for three field seasons to determine how the distribution of food and the risk of predation influenced marmots' foraging behavior. I quantified the amount of time Marmota caligata foraged in different patches of alpine meadows and assessed the distribution and abundance of vegetation eaten by marmots in these meadows. Because marmots dig burrows and run to them when attacked by predators, marmot-toburrow distance provided an index of predation risk that could be specified for different meadow patches.Patch use correlated positively with food abundance and negatively with predation risk. However, these significant relationships disappeared when partial correlations were calculated because food abundance and risk were intercorrelated. Using multiple regression, 77.0% of the variance in patch use was explained by a combination of food abundance, refuge burrow density, and a patch's distance from the talus where sleeping burrows were located. Variations in vigilance behavior (look-ups to search for predators while feeding) according to marmots' ages, the presence of other conspecifics, and animals' proximity to their sleeping burrows all indicated that predation risk influenced foraging.In a forage-manipulation experiment, the use of forage-enhanced patches increased six-fold, verifying directly the role of food availability on patch used. Concomitant with increased feeding, however, was the intense construction of refuge burrows in experimental patches that presumably reduced the risk of feeding. Thus, I suggest that food and predation risk jointly influence patch use by hoary marmots and that both factors must be considered when modeling the foraging behavior of species that can be predator and prey simultaneously.     

GPS tracking a marine predator: the effects of precision,resolution and sampling rate on foraging tracks of African Penguins

We used a prototype GPS logger to track the movements of breeding African Penguins (Spheniscus demersus). The loggers also recorded temperature and water depth, which allowed us to reconstruct foraging tracks in three dimensions, although GPS signals are interrupted when the birds dive. Here we report the loggers performance in the field and assess the effects of GPS error, resolution and sampling rate on estimates of foraging track length and speed. There is a trade-off between sampling rate and battery lifespan. We tested loggers at sampling intervals of 1 s, 10 s, 1 min, 2 min and 10 min. Sampling less frequently increases the chance of tracking an entire foraging trip, but it slows uplink times, slightly decreases the accuracy of positional fixes, and significantly reduces the ability to measure fine-scale aspects of foraging behaviour. Compared with radio or satellite tracking, GPS loggers offer unprecedented detail about animal movements. The results of our analysis suggest that techniques that sample relatively infrequently, such as satellite tracking, underestimate actual track lengths by up to 50%. However, caution is needed when interpreting fine-scale sampling for relatively slow-moving organisms. Re-sampling 1-s tracks suggests that c. 35% of apparent movements at this scale are due to measurement error and, more importantly, the limited spatial resolution of GPS (1.85×1.54 m at the study area). We recommend that researchers use a 1-s sampling rate for fine-scale studies, but resample at less frequent intervals to remove spurious noise for slow-moving animals. At current levels of resolution, animals should move at least 4 m per sampling interval. We provide empirical correction factors to compare inferred track length sampled at different rates, but caution that these are idiosyncratic and strongly dependent on the animals behaviour. Overall, GPS loggers offer a significant advance for studies of fine-scale animal movement patterns.Communicated by O. Kinne, Oldendorf/Luhe     

Male lagoon gobies, <Emphasis Type="Italic">Knipowitschia panizzae</Emphasis>, prefer more ornamented to larger females

Female ornamentation may be directly sexually selected, by male choice or female competition, or occurs as the result of a genetic correlation, arising from sexual selection on males. However, increasing evidence supports the former hypothesis, suggesting that males actively choose their partner preferring traits indicative of female quality. In the lagoon goby, Knipowitschia panizzae, a polygynous species whose males perform parental care to eggs, body length and the size of a sex-specific yellow patch on the belly are known to be reliable indicators of female fecundity. In this paper, we tested, using dummies, the male’s mating preferences for female body and yellow belly patch sizes. The two experimental trials in which a single female trait was variable showed that males prefer a larger belly patch and a larger body size, indicating that both these characters are selected by male mate choice. However, when faced with dummies exhibiting an inverse combination of body and belly patch sizes (experiment 3), males significantly preferred the smaller ones with larger yellow belly patches. A calculation of dummy theoretical fecundity reveals that in the first two experiments, males would have received an immediate benefit from their choice in terms of egg number, whereas in the third one, males chose partners that would have provided them with fewer eggs. The male lagoon goby preference for females with larger belly patches, regardless of their size, suggests that this trait, in addition to indicating fecundity, conveys information about other aspects of female and/or egg quality.     

Movement of foraging Tundra Swans explained by spatial pattern in cryptic food densities

We tested whether Tundra Swans use information on the spatial distribution of cryptic food items (below ground Sago pondweed tubers) to shape their movement paths. In a continuous environment, swans create their own food patches by digging craters, which they exploit in several feeding bouts. Series of short (<1 m) intra-patch movements alternate with longer inter-patch movements (>1 m). Tuber biomass densities showed a positive spatial auto-correlation at a short distance (<3 m), but not at a larger distance (3-8 m). Based on the spatial pattern of the food distribution (which is assumed to be pre-harvest information for the swan) and the energy costs and benefits for different food densities at various distances, we calculated the optimal length of an inter-patch movement. A swan that moves to the patch with the highest gain rate was predicted to move to the adjacent patch (at 1 m) if the food density in the current patch had been high (>25 g/m2) and to a more distant patch (at 7-8 m) if the food density in the current patch had been low (<25 g/m2). This prediction was tested by measuring the response of swans to manipulated tuber densities. In accordance with our predictions, swans moved a long distance (>3 m) from a low-density patch and a short distance (<3 m) from a high-density patch. The quantitative agreement between prediction and observation was greater for swans feeding in pairs than for solitary swans. The result of this movement strategy is that swans visit high-density patches at a higher frequency than on offer and, consequently, achieve a 38% higher long-term gain rate. Swans also take advantage of spatial variance in food abundance by regulating the time in patches, staying longer and consuming more food from rich than from poor patches. We can conclude that the shape of the foraging path is a reflection of the spatial pattern in the distribution of tuber densities and can be understood from an optimal foraging perspective.