Lisa A. Leaver

Food caching and differential cache pilferage: a field study of coexistence of sympatric kangaroo rats and pocket mice

Ecologists studying sympatric heteromyid rodents have sought evidence for species differences in primary foraging abilities and preferences and/or behavioural responses to predation risk in order to explain coexistence. The present field study was conducted to test the hypothesis that another factor may be involved, namely differences in caching patterns, which may result in differences in vulnerability to pilferage. We examined differences between kangaroo rats (Dipodomys merriami) and pocket mice (Chaetodipus spp.) in foraging, caching and pilferage behaviour. Specifically, we examined interactions at food patches, differential food caching patterns, and differential vulnerability to cache pilferage. Observations conducted at artificial seed patches showed that kangaroo rats dominated access to the patches by arriving and foraging first and by chasing pocket mice away. Individually provisioned pocket mice stored most seeds in underground burrows (larder hoarding), whereas kangaroo rats predominantly cached seeds in small, spatially dispersed caches in shallow pits in the surface of the sand (scatter hoarding). Pocket mice pilfered from each other as well as from the kangaroo rats, but the kangaroo rats rarely pilfered, and the only instance was from another kangaroo rat. Kangaroo rats and pocket mice were both vulnerable to cache pilferage. The results suggest that coexistence of kangaroo rats and pocket mice may be facilitated by a trade-off between primary harvest ability and the ability to exploit a resource that has been processed by another species, namely pilferage ability.

Audience effects on food caching in grey squirrels (Sciurus carolinensis): evidence for pilferage avoidance strategies

If food pilferage has been a reliable selection pressure on food caching animals, those animals should have evolved the ability to protect their caches from pilferers. Evidence that animals protect their caches would support the argument that pilferage has been an important adaptive challenge. We observed naturally caching Eastern grey squirrels (Sciurus carolinensis) in order to determine whether they used any evasive tactics in order to deter conspecific and heterospecific pilferage. We found that grey squirrels used evasive tactics when they had a conspecific audience, but not when they had a heterospecific (corvid) audience. When other squirrels were present, grey squirrels spaced their caches farther apart and preferentially cached when oriented with their backs to other squirrels, but no such effect was found when birds were present. Our data provide the first evidence that caching mammals are sensitive to the risk of pilferage posed by an audience of conspecifics, and that they utilise evasive tactics that should help to minimise cache loss. We discuss our results in relation to recent theory of reciprocal pilferage and compare them to behaviours shown by caching birds.

Using ecology to guide the study of cognitive and neural mechanisms of different aspects of spatial memory in food-hoarding animals

Understanding the survival value of behaviour does not tell us how the mechanisms that control this behaviour work. Nevertheless, understanding survival value can guide the study of these mechanisms. In this paper, we apply this principle to understanding the cognitive mechanisms that support cache retrieval in scatter-hoarding animals. We believe it is too simplistic to predict that all scatter-hoarding animals will outperform non-hoarding animals on all tests of spatial memory. Instead, we argue that we should look at the detailed ecology and natural history of each species. This understanding of natural history then allows us to make predictions about which aspects of spatial memory should be better in which species. We use the natural hoarding behaviour of the three best-studied groups of scatter-hoarding animals to make predictions about three aspects of their spatial memory: duration, capacity and spatial resolution, and we test these predictions against the existing literature. Having laid out how ecology and natural history can be used to predict detailed cognitive abilities, we then suggest using this approach to guide the study of the neural basis of these abilities. We believe that this complementary approach will reveal aspects of memory processing that would otherwise be difficult to discover.

Grazing and vigilance by Soay sheep on Lundy island : Influence of group size, terrain and the distribution of vegetation

Animals allocate the time spent on different behaviours according to nutritional requirements, the distribution of food and the risk of predation. When the perceived predation risk is high, animals primarily behave in ways to increase their safety such as regularly scanning their surroundings, forming large groups and feeding in habitats with cover or high visibility. This study investigated which factors Soay sheep (Ovis aries) are most sensitive to in allocating time to different behaviours when the risk of predation is negligible. Continuous focal sampling was used to record the behaviour of the free-ranging Soay sheep on Lundy Island and measures of vegetation, group size and terrain were also recorded. Stepwise multiple regression produced a model of grazing with terrain and range of grass coverage as predictors and a model of vigilance with terrain as a predictor. Sheep grazed for longer on the slopes and as the range of grass cover decreased. They were more vigilant on the slopes. The results support the view that the group size effect largely depends on anti-predatory vigilance and suggest that time-budgets are more sensitive to other factors in the absence of predation.

Effects of Food Preference on Scatter-Hoarding by Kangaroo Rats (Dipodomys Merriami)

Two laboratory studies were conducted to determine whether Merriams kangaroo rats invest greater effort in the caching of a more preferred food. As predicted, more of the preferred food was cached and yet the individual caches were smaller. The second experiment showed wider dispersion of the preferred food, and these caches were placed further away from the source. These findings imply that investment in protecting food from pilferage is adjusted in relation to the animals evaluation of that food.

Grey squirrels (Sciurus carolinensis) show a feature-negative effect specific to social learning

Previous laboratory studies on social learning suggest that some animals can learn more readily if they first observe a conspecific demonstrator perform the task unsuccessfully and so fail to obtain a food reward than if they observe a successful demonstrator that obtains the food. This effect may indicate a difference in how easily animals are able to associate different outcomes with the conspecific or could simply be the result of having food present in only some of the demonstrations. To investigate we tested a scatter-hoarding mammal, the eastern grey squirrel, on its ability to learn to choose between two pots of food after watching a conspecific remove a nut from one of them on every trial. Squirrels that were rewarded for choosing the opposite pot to the conspecific chose correctly more frequently than squirrels rewarded for choosing the same pot (a feature-negative effect). Another group of squirrels was tested on their ability to choose between the two pots when the rewarded option was indicated by a piece of card. This time, squirrels showed no significant difference in their ability to learn to choose the same or the opposite pot. The results add to anecdotal reports that grey squirrels can learn by observing a conspecific and suggest that even when all subjects are provided with demonstrations with the same content, not all learning occurs equally. Prior experience or expectations of the association between a cue (a conspecific) and food influences what can be learned through observation whilst previously unfamiliar cues (the card) can be associated more readily with any outcome.

Touch screen assays of behavioural flexibility and error characteristics in Eastern grey squirrels (Sciurus carolinensis)

Behavioural flexibility allows animals to adjust their behaviours according to changing environmental demands. Such flexibility is frequently assessed by the discrimination–reversal learning task. We examined grey squirrels’ behavioural flexibility, using a simultaneous colour discrimination–reversal learning task on a touch screen. Squirrels were trained to select their non-preferred colour in the discrimination phase, and their preferred colour was rewarded in a subsequent reversal phase. We used error rates to divide learning in each phase into three stages (perseveration, chance level and ‘learned’) and examined response inhibition and head-switching during each stage. We found consistent behavioural patterns were associated with each learning stage: in the perseveration stage, at the beginning of each training phase, squirrels showed comparable response latencies to correct and incorrect stimuli, along with a low level of head-switching. They quickly overcame perseveration, typically in one to three training blocks. In the chance-level stage, response latencies to both stimuli were low, but during initial discrimination squirrels showed more head-switches than in the previous stage. This suggests that squirrels were learning the current reward contingency by responding rapidly to a stimulus, but with increased attention to both stimuli. In the learned stage, response latencies to the correct stimulus and the number of head-switches were at their highest, whereas incorrect response latencies were at their lowest, and differed significantly from correct response latencies. These results suggest increased response inhibition and attention allowed the squirrels to minimise errors. They also suggest that errors in the ‘learned’ stage were related to impulsive emission of the pre-potent or previously learned responses.

Serial reversal learning in Gray Squirrels: learning efficiency as a function of learning and change of tactics

Learning allows individuals to adapt their behaviors flexibly to a changing environment. When the same change recurs repeatedly, acquiring relevant tactics may increase learning efficiency. We examined this relationship, along with the effects of proactive interference and other interference information, in a serial spatial reversal task with 5 gray squirrels (Sciurus carolinensis). Squirrels completed an acquisition and 11 reversal phases with a poke box in which 2 of 4 possible reward locations were baited diagonally in a square array. In this situation, an efficient tactic is to locate the diagonally related locations consecutively (integrative search tactic) instead of searching rewards in a clockwise or counterclockwise direction (sequential search tactic). All squirrels formed a learning set, acquiring successive reversals in fewer trials. Although 4 squirrels gradually employed more integrative tactics in locating the rewards both within and between phases, sequential tactics were used in the first trial of each phase. This suggests the integrative tactic did not depend on an association between the rewarded locations but was learned as a spatial pattern and/or by use of extra-apparatus cues to locate individual rewards. Generalized estimating equation models showed that learning efficiency increased with experience and tactic change. Although tactic change partially mediated the effect of learning on learning efficiency, learning remained an independent contribution to improved efficiency. Squirrels that used more integrative tactics made fewer total errors than squirrels that used less integrative tactics, suggesting that learning a task-relevant tactic using spatial cues can provide direct benefits in maximizing rewards and minimizing time costs.

Behavioral flexibility versus rules of thumb: how do grey squirrels deal with conflicting risks?

In order to test how flexibly animals are able to behave when making trade-offs that involve assessing constantly changing risks, we examined whether wild Eastern grey squirrels showed flexibility of behavioral responses in the face of variation in 2 conflicting risks, cache pilferage, and predation. We established that cache pilferage risk decreased with distance from cover and was thus negatively correlated with long-term predation risk. We then measured changes in foraging and food-caching behavior in the face of changes in the risk of predation and food theft over a short time-scale. We found that, overall, squirrels move further away from the safety of cover when they cache, compared to when they forage, as predicted by pilferage risk. However, there was no effect of immediate pilferage or predation risk (i.e., the presence of potential predators or pilferers) on the distance from cover at which they cached, and only a slight increase in forage distance when predation risk increased. These results suggest that “rules of thumb” based on static cues may be more cost-effective for assessing risk than closely tracking changes over time in the way suggested by a number of models of risk assessment.

Multiple feature use in pigeons’ category discrimination: The influence of stimulus set structure and the salience of stimulus differences

Two experiments investigated what makes it more likely that pigeons’ behavior will come under the control of multiple relevant visual stimulus dimensions. Experiment 1 investigated the effect of stimulus set structure, using a conditional discrimination between circles that differed in both hue and diameter. Two training conditions differed in whether hue and diameter were correlated in the same way within positive and negative stimulus sets as between sets. Transfer tests showed that all pigeons came under the control of both dimensions, regardless of stimulus set structure. Experiment 2 investigated the effect of the relative salience of the stimulus differences on three visual dimensions. Pigeons learned a multiple simultaneous discrimination between circular patches of sinusoidal gratings that differed in hue, orientation, and spatial frequency. In initial training, each stimulus only included one positive or negative feature, and the stimulus differences on the three dimensions were adjusted so that the rates of learning about the three dimensions were kept approximately equal. Transfer tests showed that all three dimensions acquired control over behavior, with no single dimension dominating consistently across pigeons. Subsequently the pigeons were trained in a polymorphous category discrimination using all three dimensions, and the level of control by the three dimensions tended to become more equal as polymorphous training continued. We conclude that the salience of the stimulus differences on different dimensions is an important factor in whether pigeons will come under the control of multiple dimensions of visual stimuli.