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Devenport, J. A., Patterson, M. R., & Devenport, L. D. (2005). Dynamic averaging and foraging decisions in horses (Equus callabus). J. Comp. Psychol., 119(3), 352–358.
Abstract: The variability of most environments taxes foraging decisions by increasing the uncertainty of the information available. One solution to the problem is to use dynamic averaging, as do some granivores and carnivores. Arguably, the same strategy could be useful for grazing herbivores, even though their food renews and is more homogeneously distributed. Horses (Equus callabus) were given choices between variable patches after short or long delays. When patch information was current, horses returned to the patch that was recently best, whereas those without current information matched choices to the long-term average values of the patches. These results demonstrate that a grazing species uses dynamic averaging and indicate that, like granivores and carnivores, they can use temporal weighting to optimize foraging decisions.
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Fischhoff, I. R., Sundaresan, S. R., Cordingley, J., Larkin, H. M., Sellier, M. - J., & Rubenstein, D. I. (2007). Social relationships and reproductive state influence leadership roles in movements of plains zebra, Equus burchellii. Anim. Behav., 73(5), 825–831.
Abstract: In animal groups, collective movements emerge from individual interactions. Biologists seek to identify how characteristics of actors in these groups, and their relationships, influence the decision-making process. We distinguished two basic factors determining leadership in group choices: identity and state. We hypothesized that identity is more important to leadership in groups with stable relationships, which permit the development of habitual roles. In groups with fluid membership, particular individuals or subgroups are less likely to emerge as consistent leaders. Instead, we predicted that movement initiation in unstable groups depends on individual state at the time of the decision. We characterized how identity and reproductive state influenced leadership patterns in the movements of plains zebra. As in many other mammals, lactation in this species significantly alters water and energy needs. We investigated leadership in tightly knit harems and loosely bonded herds of multiple harems. Harem females tended to have habitual roles in the initiation of harem movement. In herds, however, we found no consistent leaders among harems. At both levels of social organization, lactation was a key determinant of leadership. In harems, lactating females were more likely to initiate movement than nonlactating females. In turn, harems containing lactating females were more likely to lead herd movements. Thus, we conclude that social relationships and reproductive state together shape the interactions that produce group behaviours. One benefit to lactating females of leading herd movements is preferential access to scarce water. Thus, leadership roles in group decisions may have fitness consequences.
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Waite, T. A. (2002). Interruptions improve choice performance in gray jays: prolonged information processing versus minimization of costly errors. Anim. Cogn., 5(4), 209–214.
Abstract: Under the assumption that selection favors minimization of costly errors, erroneous choice may be common when its fitness cost is low. According to an adaptive-choice model, this cost depends on the rate at which an animal encounters the choice: the higher this rate, the smaller the cost of choosing a less valuable option. Errors should thus be more common when interruptions to foraging are shorter. A previous experiment supported this prediction: gray jays, Perisoreus canadensis, were more error prone when subjected to shorter delays to access to food rewards. This pattern, though, is also predicted by an attentional-constraints model. Because the subjects were able to inspect the rewards during delays, their improved performance when subjected to longer delays could have been a byproduct of the experimentally prolonged opportunity for information processing. To evaluate this possibility, a follow-up experiment manipulated both delay to access and whether rewards could be inspected during delays. Depriving jays of the opportunity to inspect rewards (using opaque lids) induced only a small, nonsignificant increase in error rate. This effect was independent of length of delay and so the jays' improved performance when subjected to longer delays was not simply a byproduct of prolonged information processing. More definitively, even when the jays were prevented from inspecting rewards during delays, their performance improved when subjected to longer delays. The findings are thus consistent with the adaptive-choice model.
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Tebbich, S., Bshary, R., & Grutter, A. S. (2002). Cleaner fish Labroides dimidiatus recognise familiar clients. Anim. Cogn., 5(3), 139–145.
Abstract: Individual recognition has been attributed a crucial role in the evolution of complex social systems such as helping behaviour and cooperation. A classical example for interspecific cooperation is the mutualism between the cleaner fish Labroides dimidiatus and its client reef fish species. For stable cooperation to evolve, it is generally assumed that partners interact repeatedly and remember each other's past behaviour. Repeated interactions may be achieved by site fidelity or individual recognition. However, as some cleaner fish have more than 2,300 interactions per day with various individuals per species and various species of clients, basic assumptions of cooperation theory might be violated in this mutualism. We tested the cleaner L. dimidiatus and its herbivorous client, the surgeon fish Ctenochaetus striatus, for their ability to distinguish between a familiar and an unfamiliar partner in a choice experiment. Under natural conditions, cleaners and clients have to build up their relationship, which is probably costly for both. We therefore predicted that both clients and cleaners should prefer the familiar partner in our choice experiment. We found that cleaners spent significantly more time near the familiar than the unfamiliar clients in the first 2 minutes of the experiment. This indicates the ability for individual recognition in cleaners. In contrast, the client C. striatus showed no significant preference. This could be due to a sampling artefact, possibly due to a lack of sufficient motivation. Alternatively, clients may not need to recognise their cleaners but instead remember the defined territories of L. dimidiatus to achieve repeated interactions with the same individual.
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Petherick, J. C., Seawright, E., & Waddington, D. (1993). Influence of motivational state on choice of food or a dustbathing/foraging substrate by domestic hens. Behav. Process., 28(3), 209–220.
Abstract: Domestic hens were trained to run a Y-maze and make an association between differently coloured doorways and access to food pellets or sand. The hens were tested for their choice of doorway when the goals were not visible from the choice point and when they were food or sand deprived. Hens made the choice appropriate to their deprivation state (correct choice) significantly more often for food than sand and were faster at choosing and entering the goal box when food deprived. In a follow up experiment, the goals were visible from the choice point. Again the hens chose correctly significantly more often when food than sand deprived and made the choice and entered the goal box faster when food deprived. Thus, failure to choose sand in the first experiment was not due to an inability to learn the association, but appears to result from a strong motivation to feed in the Y-maze, even when not food deprived, and a weak motivation to dustbathe or forage, even when sand deprived.
Keywords: Food; Learning; Litter; Motivation; Poultry; Preference
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Lingle, S., Rendall, D., Wilson, W. F., DeYoung, R. W., & Pellis, S. M. (2007). Altruism and recognition in the antipredator defence of deer: 2. Why mule deer help nonoffspring fawns. Anim. Behav., 73(5), 907–916.
Abstract: Both white-tailed deer, Odocoileus virginianus, and mule deer, O. hemionus, females defend fawns against coyotes, Canis latrans, but only mule deer defend nonoffspring conspecific and heterospecific fawns. During a predator attack, females may have to decide whether to defend a fawn while having imperfect information on its identity obtained from hearing a few distress calls. Although imperfect recognition can influence altruistic behaviour, few empirical studies have considered this point when testing functional explanations for altruism. We designed a series of playback experiments with fawn distress calls to test alternative hypotheses (by-product of parental care, kin selection, reciprocal altruism) for the mule deer's defence of nonoffspring, specifically allowing for the possibility that females mistake these fawns for their own. White-tailed deer females approached the speaker only when distress calls of white-tailed deer fawns were played and when their own fawn was hidden, suggesting that fawn defence was strictly a matter of parental care in this species. In contrast, mule deer females responded similarly and strongly, regardless of the caller's identity, the female's reproductive state (mother or nonmother) or the presence of their own offspring. The failure of mule deer females to adjust their responses to these conditions suggests that they do not defend nonoffspring because they mistake them for their own fawns. The lack of behavioural discrimination also suggests that kin selection, reciprocal altruism and defence of the offspring's area are unlikely to explain the mule deer's defence of nonoffspring. We identify causal and functional questions that still need to be addressed to understand why mule deer defend fawns so indiscriminately.
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Schwab, C., & Huber, L. (2006). Obey or not obey? Dogs (Canis familiaris) behave differently in response to attentional states of their owners. J Comp Psychol, 120(3), 169–175.
Abstract: Sixteen domestic dogs (Canis familiaris) were tested in a familiar context in a series of 1-min trials on how well they obeyed after being told by their owner to lie down. Food was used in 1/3 of all trials, and during the trial the owner engaged in 1 of 5 activities. The dogs behaved differently depending on the owner's attention to them. When being watched by the owner, the dogs stayed lying down most often and/or for the longest time compared with when the owner read a book, watched TV, turned his or her back on them, or left the room. These results indicate that the dogs sensed the attentional state of their owners by judging observable behavioral cues such as eye contact and eye, head, and body orientation.
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Hostetter, A. B., Cantero, M., & Hopkins, W. D. (2001). Differential use of vocal and gestural communication by chimpanzees (Pan troglodytes) in response to the attentional status of a human (Homo sapiens). J. Comp. Psychol., 115(4), 337–343.
Abstract: This study examined the communicative behavior of 49 captive chimpanzees (Pan troglodytes), particularly their use of vocalizations, manual gestures, and other auditory- or tactile-based behaviors as a means of gaining an inattentive audience's attention. A human (Homo sapiens) experimenter held a banana while oriented either toward or away from the chimpanzee. The chimpanzees' behavior was recorded for 60 s. Chimpanzees emitted vocalizations faster and were more likely to produce vocalizations as their 1st communicative behavior when a human was oriented away from them. Chimpanzees used manual gestures more frequently and faster when the human was oriented toward them. These results replicate the findings of earlier studies on chimpanzee gestural communication and provide new information about the intentional and functional use of their vocalizations.
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Søndergaard, E., Jensen, M. B., & Nicol, C. J. (). Motivation for social contact in horses measured by operant conditioning. Appl. Anim. Behav. Sci., In Press, Corrected Proof.
Abstract: Although horses are social animals they are often housed individually with limited social contact to other horses and this may compromise their welfare. The present study included eight young female horses and investigated the strength of motivation for access to full social contact, head contact and muzzle contact, respectively, to a familiar companion horse. Horses were housed individually next to their companion horse and separations between pens prevented physical contact. During daily test sessions horses were brought to a test area where they could access an arena allowing social contact. Arena access during 3 min was given after completion of a predetermined number of responses on a panel. Fixed ratios (FR) of 8, 16, 24, 32 and 40 responses per arena access were applied in a random order, one per daily test session, within each test week (Monday to Friday), and the number of rewards per daily test session was recorded. All horses could access all three types of social contact in a cross-over design, and an empty arena was used as control. Motivational strength was assessed using elasticity of demand functions, which were estimated based on the number of rewards earned and FR. Elasticities of demand for the three types of social contact were low (-0.20), and not significantly different, although increasing FR still resulted in a decrease in rewards obtained for all three types of social contact (P < 0.001). Across FR-levels horses earned more rewards for social contact than for an empty arena, as shown by much higher intercept values (2.51 vs. 0.99; P < 0.001). However, the elasticity of demand for infrequent access to an empty arena (-0.08) was lower than for social contact (P < 0.01) and not significantly different from zero (P = 0.07). Horses performed more social behaviour the lesser the restriction on social contact (full > head > muzzle). However, the finding that horses showed a similar and high motivation for all three types of social contact suggests that they are valued equally highly in a situation where the alternative is no social contact.
Keywords: Horse; Housing; Social behaviour; Operant conditioning; Motivation
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Søndergaard, E., Jensen, M. B., & Nicol, C. J. (2011). Motivation for social contact in horses measured by operant conditioning. Appl. Anim. Behav. Sci., 132(3-4), 131–137.
Abstract: Although horses are social animals they are often housed individually with limited social contact to other horses and this may compromise their welfare. The present study included eight young female horses and investigated the strength of motivation for access to full social contact, head contact and muzzle contact, respectively, to a familiar companion horse. Horses were housed individually next to their companion horse and separations between pens prevented physical contact. During daily test sessions horses were brought to a test area where they could access an arena allowing social contact. Arena access during 3 min was given after completion of a predetermined number of responses on a panel. Fixed ratios (FR) of 8, 16, 24, 32 and 40 responses per arena access were applied in a random order, one per daily test session, within each test week (Monday to Friday), and the number of rewards per daily test session was recorded. All horses could access all three types of social contact in a cross-over design, and an empty arena was used as control. Motivational strength was assessed using elasticity of demand functions, which were estimated based on the number of rewards earned and FR. Elasticities of demand for the three types of social contact were low (-0.20), and not significantly different, although increasing FR still resulted in a decrease in rewards obtained for all three types of social contact (P < 0.001). Across FR-levels horses earned more rewards for social contact than for an empty arena, as shown by much higher intercept values (2.51 vs. 0.99; P < 0.001). However, the elasticity of demand for infrequent access to an empty arena (-0.08) was lower than for social contact (P < 0.01) and not significantly different from zero (P = 0.07). Horses performed more social behaviour the lesser the restriction on social contact (full > head > muzzle). However, the finding that horses showed a similar and high motivation for all three types of social contact suggests that they are valued equally highly in a situation where the alternative is no social contact.
Keywords: Horse; Housing; Social behaviour; Operant conditioning; Motivation
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