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Fawcett, T. W., Skinner, A. M. J., & Goldsmith, A. R. (2002). A test of imitative learning in starlings using a two-action method with an enhanced ghost control. Anim. Behav., 64(4), 547–556.
Abstract: Imitative learning, in which an individual learns to reproduce the behaviour pattern of another, has attracted considerable attention as a potentially powerful form of social learning. Despite extensive research, however, it has proved difficult to demonstrate in nonhuman animals. We investigated the ability of European starlings, Sturnus vulgaris, to imitate the behaviour of a conspecific. Subjects watched a trained conspecific manipulating a plug for access to a food reward, using either a pushing or a pulling action. When later tested with the same apparatus these birds completed the task using the same action they had previously observed. In a second experiment, a separate group of starlings saw the plug move upwards or downwards automatically and a nearby conspecific obtain a food reward. When given access to the task these starlings failed to move the plug in the direction they had seen. Our experiment is an improvement on previous bidirectional control designs and provides strong evidence that starlings are capable of imitation. We advocate further use of this experimental design in attempts to demonstrate imitative learning. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.
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Kenward, B., Rutz, C., Weir, A. A. S., & Kacelnik, A. (2006). Development of tool use in New Caledonian crows: inherited action patterns and social influences. Anim. Behav., 72(6), 1329–1343.
Abstract: New Caledonian crows, Corvus moneduloides, are the most advanced avian tool makers and tool users. We previously reported that captive-bred isolated New Caledonian crows spontaneously use twig tools and cut tools out of Pandanus spp. tree leaves, an activity possibly under cultural influence in the wild. However, what aspects of these behaviours are inherited and how they interact with individual and social experience remained unknown. To examine the interaction between inherited traits, individual learning and social transmission, we observed the ontogeny of twig tool use in hand-reared juveniles. Successful food retrieval was preceded by stereotyped object manipulation action patterns that resembled components of the mature behaviour, demonstrating that tool-oriented behaviours in this species are an evolved specialization. However, there was also an effect of social learning: juveniles that had received demonstrations of twig tool use by their human foster parent showed higher levels of handling and insertion of twigs than did their naive counterparts; a choice experiment showed that they preferred to handle objects that they had seen being manipulated by their human foster parent. Our observations are consistent with the hypothesis that individual learning, cultural transmission and creative problem solving all contribute to the acquisition of the tool-oriented behaviours in the wild, but inherited species-typical action patterns have a greater role than has been recognized.
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Pompilio, L., & Kacelnik, A. (2005). State-dependent learning and suboptimal choice: when starlings prefer long over short delays to food. Anim. Behav., 70(3), 571–578.
Abstract: Recent studies have used labels such as `work ethics', `sunk costs' and `state-dependent preferences' for apparent anomalies in animals' choices. They suggest that preference between options relates to the options' history, rather than depending exclusively on the expected payoffs. For instance, European starlings, Sturnus vulgaris, trained to obtain identical food rewards from two sources while in two levels of hunger preferred the food source previously associated with higher hunger, regardless of the birds' state at the time of testing. We extended this experimentally and theoretically by studying starlings choosing between sources that differed not only in history but also in the objective properties (delay until reward) of the payoffs they delivered. Two options (PF and H) were initially presented in single-option sessions when subjects were, respectively, prefed or hungry. While option PF offered a delay until reward of 10 s in all treatments, option H delivered delays of 10, 12.5, 15 and 17.5 s in four treatments. When training was completed, we tested preference between the options. When delays in both options were equal (10 s), the birds strongly preferred H. When delay in H was 17.5 s, the birds were indifferent, with intermediate results for intermediate treatments. Preference was not mediated by disrupted knowledge of the delays. Thus, preferences were driven by past state-dependent gains, rather than by the joint effect of the birds' state at the time of choice and knowledge of the absolute properties of each alternative, as assumed in state-dependent, path-independent models of optimal choice.
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Fernández-Juricic, E., Smith, R., & Kacelnik, A. (2005). Increasing the costs of conspecific scanning in socially foraging starlings affects vigilance and foraging behaviour. Anim. Behav., 69(1), 73–81.
Abstract: Social foragers receive and use information both about companions (social information) and about events external to the group, such as presence of potential predators. Scanning behaviour is often incorporated in theoretical models using simplifying assumptions in relation to the trade-off in information gathering between body postures (head-up versus head-down); however, some avian visual systems may allow individuals to scan in both body postures. We studied these issues experimentally, using starlings, Sturnus vulgaris, foraging in enclosures on natural fields. We varied the availability of information from conspecifics by placing visual barriers that blocked the view when the subjects were in head-down position and by manipulating the distance between group members. We found that as social information was reduced, starlings spent more time scanning (on and off the ground) and head-up scanning was mainly oriented towards conspecifics. The visual-obstruction effects imply that some information about conspecifics is normally gathered while starlings are foraging head-down. Neighbour distance and visual obstruction negatively affected food-searching rates and intake rates in two ways: (1) the effect of obstruction was mediated mostly through time competition between foraging and scanning on the ground, and (2) the effect of distance was due to a reduction in the rate of prey returns per searching effort while the birds were head-down. We conclude that the head-up posture is only one component of scanning, that the effects of head-down scanning should also be considered in species with ample visual fields, and that scanning in starlings is strongly connected to monitoring other flock members.
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Schuck-Paim, C., & Kacelnik, A. (2002). Rationality in risk-sensitive foraging choices by starlings. Anim. Behav., 64(6), 869–879.
Abstract: Normative models of choice usually predict preferences between alternatives by computing their value according to some criterion, then identifying the alternative with greatest value. An important consequence of this procedure is captured in the economic concept of rationality, defined through a number of principles that are necessary for the existence of an ordinal scale of value upon which organisms base their choices. Violations of these principles, such as some recently reported breaches of transitivity and regularity in birds and honeybees, have strong implications for the understanding of decision mechanisms in humans and nonhumans alike. We investigated rationality in risky choice using European starlings, Sturnus vulgaris. Birds had to choose between two or three food sources, each associated with a different variance in delay to reward. In three experiments, starlings were strongly risk prone, showing regular and consistent preferences in binary and trinary choices. Preferences also satisfied weak and strong stochastic transitivity. Our results extend the generality of previous research in risk-sensitive foraging to situations where more than two alternatives are present and suggest that violations of rationality in risk-sensitive choices may be expressed only under restricted sets of conditions. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.
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Kacelnik, A., & Marsh, B. (2002). Cost can increase preference in starlings. Anim. Behav., 63(2), 245–250.
Abstract: We used European starlings, Sturnus vulgaris, to investigate the relationship between the cost paid to obtain food rewards and preference between stimuli associated with the resulting rewards. In no-choice trials either 16 1-m flights (high effort) or four 1-m flights (low effort) gave access to differently coloured keys. Pecking at these keys resulted in identical food rewards. When subjects were given choices between the coloured keys in choice trials without having paid any effort, the majority preferred the coloured key that was paired with the higher level of work in no-choice trials. We relate our findings to results in animal behaviour, psychology and economics, and give a theoretical account that has implications for phenomena variously recognized as the `sunk cost fallacy' (the tendency to invest more in something after much has already been invested), `work ethics' (valuing an option more as a result of physical effort), `cognitive dissonance' (making mental effort to overlook or re-evaluate information that does not accord with a dominant internal representation) and the `Concorde Fallacy' (the readiness to forego more fitness for something that has been responsible for greater fitness compromise in the past).
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Bateson, M., & Kacelnik, A. (1997). Starlings' preferences for predictable and unpredictable delays to food. Anim. Behav., 53(6), 1129–1142.
Abstract: Risk-sensitive foraging theory is based on the premise that unpredictable runs of good or bad luck can cause a variable food source to differ in fitness value from a fixed food source yielding the same average rate of gain but no unpredictability. Thus, risk-sensitive predictions are dependent on the food intake from variable sources being not only variable but also unpredictable or `risky' in outcome. This study tested whether unpredictability is a component of the value that foraging starlings,Sturnus vulgarisattribute to food sources that are variable in the delay to obtain food. Two groups of birds chose between a fixed and a variable delay option; the variable option was unpredictable in the risky group and predictable in the risk-free group in the overall rate of intake it yielded. In both groups the fixed option was adjusted by titration to quantify the magnitude of preference for predictable and unpredictable variance. On negative energy budgets both groups were significantly risk-prone, with the risky group being significantly more risk-prone than the risk-free group. Switching the birds to positive budgets by doubling the size of each food reward had no significant effect on preference, and similar trends to those found with negative budgets were observed. These results are not readily explained by risk-sensitive foraging theory, but may be explained by the algorithm used by the birds to attribute value to average expected rewards.
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Brunner, D., Kacelnik, A., & Gibbon, J. (1996). Memory for inter-reinforcement interval variability and patch departure decisions in the starling,Sturnus vulgaris. Anim. Behav., 51(5), 1025–1045.
Abstract: An experiment with starlings was conducted to investigate the effect of variability in inter-reinforcement intervals on foraging decisions. The experimental design simulated an environment in which food was distributed in patches. Patches contained zero to four food items which could be collected by pecking at a key. All patches ended with sudden depletion. The time elapsed since the last reinforcement was the only way to detect the depletion of the patch. Once a patch was depleted, a new patch could be reached by completion of a travel requirement of 20 flights between two perches. Key pecks within a patch and the time of the last response in a patch (giving-in time) were recorded. The level of variability in the inter-reinforcement intervals was varied between different conditions. An increase in inter-reinforcement interval variability resulted in a flattening of response rate functions and giving-in time distributions, and in more asymmetry of the response functions, but not of the giving-in time distributions. Two theoretical models of decision making are presented, which differ in the assumptions about memory constraints. In one case, all inter-reinforcement intervals are remembered but in the other, only the intervals with extreme values are remembered. Both models accommodate response rates as a function of trial time, but only the second is compatible with the observed departure decision. Our results are compatible with net rate maximization.
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Bateson, M., & Kacelnik, A. (1995). Accuracy of memory for amount in the foraging starling,Sturnus vulgaris. Anim. Behav., 50(2), 431–443.
Abstract: Attempts to include psychological constraints in models of foraging behaviour differ in their assumptions concerning the accuracy of estimation of environmental parameters. Psychologists model estimation error as increasing linearly with the magnitude of a stimulus (Weber's Law), whereas behavioural ecologists either ignore error or assume it to be independent of stimulus magnitude. Studies on the estimation of time intervals have confirmed Weber's Law, but there are few data on the accuracy of estimation of amounts of food. Since the currency of most foraging models is the amount of food acquired per unit of time spent foraging, information on estimation of amount is required. Here, a titration method was used in which starlings chose between two cues. One colour signalled a standard food reward, and the other a reward that adjusted in magnitude according to the birds' choices: it increased when the standard was preferred and decreased when the adjusting option was preferred. There were two standards of 3 and 9 units of food, each of which was delivered at two rates to control for possible effects of rate of reinforcement on discrimination. The observed value of the adjusting option oscillated around a mean value slightly larger than that of the standard. The amplitude and period of these oscillations were larger when the standard was larger, independent of the rate of reinforcement. Also, molecular analysis showed that the probability of choosing the currently larger alternative increased as the relative difference between the adjusting option and standard increased. These results are consistent with Weber's Law applying to starlings' memories for amounts of food.
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Todd, I. A., & Kacelnik, A. (1993). Psychological mechanisms and the Marginal Value Theorem: dynamics of scalar memory for travel time. Anim. Behav., 46(4), 765–775.
Abstract: Abstract. The relation between memory for travel time and foraging decisions was studied experimentally. The temporal properties of two environments with patchily distributed food were simulated in the laboratory using pigeons, Columba livia, as subjects. The two environments differed in mean travel time, while the coefficient of variation of travel time and the decelerated function relating cumulative food gain to time in the patch were held constant within and between environments. Each environment contained a uniform mixture of five travel times experienced in a random order. Two of the five travel times were common in both environments. Effects of travel time were studied by comparing prey collected per patch visit (PPV) after various travel times within each environment, and by comparing patch exploitation after equal travel times between environments. Within the environment with long mean travel time (LMT) PPV was positively correlated with the last and the penultimate travel times but not with travel times before that. The increase in PPV per second of last travel time was six times greater than the increase per second of penultimate travel time, implying very steep memory discounting. In the environment with short mean travel time (SMT), there was no correlation between PPV and previous travel times. However, comparisons between environments of visits following travel times common to both environments (thus removing the effect of the last travel time) showed that substantially more prey were taken after equal travel times in the LMT than in the SMT environment. This difference cannot be accounted for by the within-environment effect of penultimate travel time, implying that there is a different, less steeply devalued, effect of the mixture of travel times. A model of information processing based on combining Scalar Expectancy Theory with the predictions of rate maximization under the Marginal Value Theorem is presented. The model can approximate the results obtained in this and previous experiments and provides a framework for further analysis of memory mechanisms of foraging behaviour.
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