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Held, S., Baumgartner, J., Kilbride, A., Byrne, R. W., & Mendl, M. (2005). Foraging behaviour in domestic pigs (Sus scrofa): remembering and prioritizing food sites of different value. Anim. Cogn., 8(2), 114–121.
Abstract: This experiment investigated whether domestic pigs can remember the locations of food sites of different relative value, and how a restricted retrieval choice affects their foraging behaviour. Nine juvenile female pigs were trained to relocate two food sites out of a possible eight in a spatial memory task. The two baited sites contained different amounts of food and an obstacle was added to the smaller amount to increase handling time. On each trial, a pig searched for the two baited sites (search visit). Once it had found and eaten the bait, it returned for a second (relocation) visit, in which the two same sites were baited. Baited sites were changed between trials. All subjects learnt the task. When allowed to retrieve both baits, the subjects showed no preference for retrieving a particular one first (experiment 1). When they were allowed to retrieve only one bait, a significant overall preference for retrieving the larger amount emerged across subjects (experiment 2). To test whether this preference reflected an avoidance of the obstacle with the smaller bait, 15 choice-restricted control trials were conducted. In control trials obstacles were present with both baits. Pigs continued to retrieve the larger bait, indicating they had discriminated between the two food sites on the basis of quantity or profitability and adjusted their behaviour accordingly when the relocation choice was restricted. This suggests for the first time that domestic pigs have the ability to discriminate between food sites of different relative value and to remember their respective locations.
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Krueger, K., Trager, L., Farmer, K., & Byrne, R. (2022). Tool Use in Horses. Animals, 12(15), 1876.
Abstract: Tool use has not yet been confirmed in horses, mules or donkeys. As this subject is difficult to research with conventional methods, we used a crowdsourcing approach to gather data. We contacted equid owners and carers and asked them to report and video examples of �unusual� behaviour via a dedicated website. We also searched YouTube and Facebook for videos of equids showing tool use. From 635 reports, including 1014 behaviours, we found 20 cases of tool use, 13 of which were unambiguous in that it was clear that the behaviour was not trained, caused by reduced welfare, incidental or accidental. We then assessed (a) the effect of management conditions on tool use and (b) whether the animals used tools alone, or socially, involving other equids or humans. We found that management restrictions were associated with corresponding tool use in 12 of the 13 cases (p = 0.01), e.g., equids using sticks to scrape hay within reach when feed was restricted. Furthermore, 8 of the 13 cases involved other equids or humans, such as horses using brushes to groom others. The most frequent tool use was for foraging, with seven examples, tool use for social purposes was seen in four cases, and there was just one case of tool use for escape. There was just one case of tool use for comfort, and in this instance, there were no management restrictions. Equids therefore can develop tool use, especially when management conditions are restricted, but it is a rare occurrence.
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Byrne, R. W. (2007). Culture in great apes: using intricate complexity in feeding skills to trace the evolutionary origin of human technical prowess. Phil. Trans. Biol. Sci., 362(1480), 577–585.
Abstract: Geographical cataloguing of traits, as used in human ethnography, has led to the description of “culture” in some non-human great apes. Culture, in these terms, is detected as a pattern of local ignorance resulting from environmental constraints on knowledge transmission. However, in many cases, the geographical variations may alternatively be explained by ecology. Social transmission of information can reliably be identified in many other animal species, by experiment or distinctive patterns in distribution; but the excitement of detecting culture in great apes derives from the possibility of understanding the evolution of cumulative technological culture in humans. Given this interest, I argue that great ape research should concentrate on technically complex behaviour patterns that are ubiquitous within a local population; in these cases, a wholly non-social ontogeny is highly unlikely. From this perspective, cultural transmission has an important role in the elaborate feeding skills of all species of great ape, in conveying the “gist” or organization of skills. In contrast, social learning is unlikely to be responsible for local stylistic differences, which are apt to reflect sensitive adaptations to ecology.
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Bates, L. A., Sayialel, K. N., Njiraini, N. W., Poole, J. H., Moss, C. J., & Byrne, R. W. (2008). African elephants have expectations about the locations of out-of-sight family members. Biol Lett, 4(1), 34–36.
Abstract: Monitoring the location of conspecifics may be important to social mammals. Here, we use an expectancy-violation paradigm to test the ability of African elephants (Loxodonta africana) to keep track of their social companions from olfactory cues. We presented elephants with samples of earth mixed with urine from female conspecifics that were either kin or unrelated to them, and either unexpected or highly predictable at that location. From behavioural measurements of the elephants' reactions, we show that African elephants can recognize up to 17 females and possibly up to 30 family members from cues present in the urine-earth mix, and that they keep track of the location of these individuals in relation to themselves.
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Byrne, R. W., & Russon, A. E. (1998). Learning by imitation: a hierachical approach. Behav. Brain Sci., 21, 667–721.
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Byrne, R. W. (1993). Do larger brains mean greater intelligence? Behav. Brain Sci., 16(4), 696–697.
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Byrne, R. W., Corp, N., & Byrne, J. M. (2001). Manual dexterity in the gorilla: bimanual and digit role differentiation in a natural task. Anim. Cogn., 4(3), 347–361.
Abstract: The manipulative actions of mountain gorillas Gorilla g. beringei were examined in the context of foraging on hard-to-process plant foods in the field, in particular those used in tackling thistle Carduus nyassanus. A repertoire of 72 functionally distinct manipulative actions was recorded. Many of these actions were used in several variants of grip, finger(s) and movement path, both by different individuals and by the same individual at different times. The repertoire appears somewhat greater than that observed in comparable studies of monkeys, but a far more striking difference is found in the use of differentiated actions in concert. Mountain gorillas routinely and frequently deal with problems that involve: (1) bimanual role differentiation, with the two hands taking different roles but synchronized in time and space, and (2) digit role differentiation, with independent control of parts of the same hand used for separate purposes at the same time. The independent control that allows these abilities, so crucial to human manual constructional ability, is apparently general in African great apes. Role differentiation, between and within the hand, is evidently a primitive characteristic in the human arsenal of skills.
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Stokes, E., & Byrne, R. (2001). Cognitive capacities for behavioural flexibility in wild chimpanzees (Pan troglodytes): the effect of snare injury on complex manual food processing. Anim. Cogn., 4(1), 11–28.
Abstract: In chimpanzees, it is only in the restricted context of tool use that manual and cognitive skills have been described, comparable to those that gorillas and orang-utans display in obtaining plant foods. We report the complex food preparation skills used to eat, without tools, the leaves of the tree Broussonettia papyrifera in the Sonso community of chimpanzees at Budongo Forest, Uganda. Able-bodied individuals used multi-stage techniques that required bimanual role differentiation at several stages, and were hierarchical in organisation. A total repertoire of 14 techniques was found, with strong preference in all individuals for either of two of these; 6 additional techniques were found when flowers and leaves were eaten together. However, in this community over 20% of individuals suffer from some form of upper- or lower-limb injury as a result of snares. We investigated the manner of compensation for upper-limb injury. Only the most severely injured showed reduced feeding efficiency. Injured individuals were found to use the same repertoire of techniques as able-bodied chimpanzees. We found no evidence to suggest that injured individuals were able to develop wholly novel techniques optimal for their specific injuries, although shifts in preference for particular techniques did occur. Rather, injured individuals used novel ways of achieving particular steps in the process; by “working around” their impairments; in this way, they managed to use the same techniques as the able-bodied. Since snare injuries generally befall young animals, these results suggest that chimpanzees learn techniques partly through observational learning (of, necessarily, able-bodied individuals).
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Byrne, R. W. (1999). Imitation without intentionality. Using string parsing to copy the organization of behaviour. Anim. Cogn., 2(2), 63–72.
Abstract: A theory of imitation is proposed, string parsing, which separates the copying of behavioural organization by observation from an understanding of the cause of its effectiveness. In string parsing, recurring patterns in the visible stream of behaviour are detected and used to build a statistical sketch of the underlying hierarchical structure. This statistical sketch may in turn aid the subsequent comprehension of cause and effect. Three cases of social learning of relatively complex skills are examined, as potential cases of imitation by string parsing. Understanding the basic requirements for successful string parsing helps to resolve the conflict between mainly negative reports of imitation in experiments and more positive evidence from natural conditions. Since string parsing does not depend on comprehension of the intentions of other agents or the everyday physics of objects, separate tests of these abilities are needed even in animals shown to learn by imitation.
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Hobaiter, C., & Byrne, R. (2011). Serial gesturing by wild chimpanzees: its nature and function for communication. Anim. Cogn., 14(6), 827–838.
Abstract: Chimpanzees at Budongo, Uganda, regularly gesture in series, including ‘bouts’ of gesturing that include response waiting and ‘sequences’ of rapid-fire gesturing without pauses. We examined the distribution and correlates of 723 sequences and 504 bouts for clues to the function of multigesture series. Gesturing by older chimpanzees was more likely to be successful, but the success rate of any particular gesture did not vary with signaller age. Rather, older individuals were more likely to choose successful gestures, and these highly successful gestures were more often used singly. These patterns explain why bouts were recorded most in younger animals, whereas older chimpanzees relied more on single gestures: bouts are best interpreted as a consequence of persistence in the face of failure. When at least one gesture of a successful type occurred in a sequence, that sequence was more likely to be successful; overall, however, sequences were less successful than single gestures. We suggest that young chimpanzees use sequences as a ‘fail-safe’ strategy: because they have the innate potential to produce a large and redundant repertoire of gestures but lack knowledge of which of them would be most efficient. Using sequences increases the chance of giving one effective gesture and also allows users to learn the most effective types. As they do so, they need to use sequences less; sequences may remain important for subtle interpersonal adjustment, especially in play. This ‘Repertoire Tuning’ hypothesis explains a number of results previously reported from chimpanzee gesturing.
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