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Author |
Krueger, K. |
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Title  |
Social learning and innovative behaviour in horses |
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Conference Article |
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Year |
2015 |
Publication |
Proceedings of the 3. International Equine Science Meeting |
Abbreviated Journal |
Proc. 3. Int. Equine. Sci. Mtg |
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social learning, innovative behaviour, Equus caballus, cognitive capacities |
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The evaluation of important parameters for measuring the horses’ cognitive capacities is one of the central topics of the equine behaviour team at Nürtingen-Geislingen University. Social complexity has been said to be one of the settings in which needs for cognitive capacities arise in animals. A variety of studies throughout the last two decades proved the horses’ social complexity to be far more elaborate than previously assumed. Horses form social bonds for the protection of offspring, intervene in encounters of others, identify group mates individually and easily orientate in a fission fusion society.
In such socially complex societies, animals will benefit from learning socially. In many bird and primate species the degree of social complexity correlates nicely with the species abilities for social learning. Social learning was, therefore, argued to be an indicator for elaborate mental capacities in animals. We were delighted to prove that horses actually copy social behaviour and techniques for operating a feeding apparatus from older and higher ranking group members. In a recent study we found young horses, at the age of 3 to 12, to copy the operation of a feeding apparatus from a human demonstrator. Social learning seems to work nicely in horses when the social background of the animals is considered.
The degree to which individual animals adapt to changes in their social or physical environment by finding innovative solution appears to be the other side of the coin, of whether animals adjust to challenges by social learning. It is not very astonishing, that along with the animals’ social complexity and their ability to learn socially also the degree to which they show innovative behaviour was claimed to be one of the most important demonstrations of advanced cognitive capacities. In a recent approach, we started to ask horse owners and horse keepers in many countries to tell us about unusual behaviour of their horses via a web site (http://innovative-behaviour.org). To date, we received 204 cases of innovative behaviour descriptions from which six cases were clear examples of tool use or borderline tool use. We categorized the innovative behaviours into the classes, a) innovations to gain food, b) innovations to gain freedom, c) social innovations, d) innovations to increase maintenance, and e) innovations that could not be clearly assigned to a category. About 20% of the innovative horses showed more than one innovation. These animals could be termed “true innovators”. Again, young horses were more innovative than older ones with the age group 5 – 9 showing the highest number of innovative behaviour descriptions.
In a nutshell, the horses’ cognitive capacities appear to be underestimated throughout the last decades. The horses’ social complexity is far more elaborate than previously assumed, horses learn socially from conspecific and humans, some of them demonstrate innovative behaviour adaptations to their environment and even simple forms of tool use. |
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Krueger, K. |
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Xenophon Publishing |
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Wald |
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in prep |
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978-3-95625-000-2 |
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Equine Behaviour @ team @ |
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5848 |
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Author |
McComb, K. |
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Title |
Social cognition and emotional awareness: studies on elephants and horses |
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Conference Article |
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2015 |
Publication |
Proceedings of the 3. International Equine Science Meeting |
Abbreviated Journal |
Proc. 3. Int. Equine. Sci. Mtg |
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Corporate Author |
McComb, K. |
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Xenophon Publishing |
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Wald |
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Krueger, K. |
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Proc. 3. Int. Equine. Sci. Mtg |
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in prep |
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978-3-95625-000-2 |
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Equine Behaviour @ team @ |
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5800 |
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Author |
Seyfarth, R.M.; Cheney, D.L. |
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Title |
Social cognition |
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Journal Article |
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2015 |
Publication |
Animal Behaviour |
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103 |
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191-202 |
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Keywords |
evolution; fitness; future research; personality; selective pressure; skill; social cognition |
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The social intelligence hypothesis argues that competition and cooperation among individuals have shaped the evolution of cognition in animals. What do we mean by social cognition? Here we suggest that the building blocks of social cognition are a suite of skills, ordered roughly according to the cognitive demands they place upon individuals. These skills allow an animal to recognize others by various means; to recognize and remember other animals' relationships; and, perhaps, to attribute mental states to them. Some skills are elementary and virtually ubiquitous in the animal kingdom; others are more limited in their taxonomic distribution. We treat these skills as the targets of selection, and assume that more complex levels of social cognition evolve only when simpler methods are inadequate. As a result, more complex levels of social cognition indicate greater selective pressures in the past. The presence of each skill can be tested directly through field observations and experiments. In addition, the same methods that have been used to compare social cognition across species can also be used to measure individual differences within species and to test the hypothesis that individual differences in social cognition are linked to differences in reproductive success. |
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0003-3472 |
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Equine Behaviour @ team @ |
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6025 |
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Author |
Kaplan, G. |
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Title |
Social animals and Communication, with special reference to horses |
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Conference Article |
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Year |
2015 |
Publication |
Proceedings of the 3. International Equine Science Meeting |
Abbreviated Journal |
Proc. 3. Int. Equine. Sci. Mtg |
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Corporate Author |
Kaplan, G. |
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Xenophon Publishing |
Place of Publication |
Wald |
Editor |
Krueger, K. |
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Proc. 3. Int. Equine. Sci. Mtg |
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in prep |
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978-3-95625-000-2 |
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Equine Behaviour @ team @ |
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5796 |
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Author |
Gabor, V.; Gerken, M. |
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Title |
Shetland ponies (Equus caballus) show quantity discrimination |
Type |
Conference Article |
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Year |
2015 |
Publication |
Proceedings of the 3. International Equine Science Meeting |
Abbreviated Journal |
Proc. 3. Int. Equine. Sci. Mtg |
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Cognition, numerical capacity, numerosity judgment, Shetland ponys |
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The complex housing environment and the close contact between humans and horses in equine sports place high demands on the learning capacity of horses. To date only limited information is available on the learning ability of the horse including higher order cognition. A type of higher order cognition is to perceive and discriminate quantities. Several mammals and birds have shown to be capable of discriminating objects due to their quantity (Brannon and Roitman, 2003). With regard to horses, there are only few studies available concerning their numerosity judgment (Uller and Lewis, 2009) and this ability is discussed controversially (Henselek et al, 2012). Possibly, the legacy of ‘‘Clever Hans’’ overshadowed further research on numerical capacity in horses, a horse to whom several psychologists incorrectly attributed the capacity of symbolic calculation (Pfungst, 1907; Rosenthal, 1965). In the present study we wanted to show whether Shetland ponies are able to transfer a previously learned concept of sameness to a numerosity judgment. The base of the test design was a “matching to sample” task, where the ponies had learned to relate abstract symbols to another which were presented on a LCD screen. Three Shetland ponies, which previously solved the matching to sample task, were tested in two test phases. In the first test phase different quantities of dots were presented (1 vs. 2; 2 vs. 3; 3 vs. 4; 4 vs. 5). To exclude discrimination due to the shape of the stimuli, the dots were varied in size and arrangement. The stimuli were presented in a triangular arrangement on the LCD screen; the sample stimulus was presented in the middle above and the discrimination stimuli in the two lower corners (S+ and S-). The pony received a food reward, by choosing the positive stimulus (S+). When the negative stimulus (S-) was chosen, the pony entered the next trial. Each learning session consisted of 20 decision trials. To investigate whether the numerosity judgment was transferable to mixed geometrical symbols (tri-, rectangle, rhomb, dot and cross) a second test phase was designed. All of the three Shetland ponies met the learning criterion of the first test phase (80% correct responses in two consecutive sessions) within the first eight sessions. One pony could transfer all judgments to the mixed symbols (up to 4 vs. 5), another pony up to 3 vs. 5 and the third on the level 2 vs. 3. These are the first reported findings that ponies are able to discriminate up to five objects. The numerosity judgment seemed to be easier for the ponies when homogenous objects were presented, than in the case of heterogeneous symbols. The reaction of the ponies occurred within few seconds, suggesting that the animals used subitizing for their numerosity judgment.
Keywords:
Cognition, numerical capacity, numerosity judgment, Shetland ponys
References:
Brannon E, Roitman J (2003) Nonverbal representations of time and number in animals and human infants. In: Meck WH (ed) Functional and neural mechanisms of interval timing. CRC Press, Boca Raton, pp 143–182
Henselek Y, Fischer J, Schloegl C (2012) Does the stimulus type influence horses’ performance in a quantity discrimination task? Front Psychol 3:504
Pfungst O (1907) Der Kluge Hans. Ein Beitrag zur nicht-verbalen Kommunikation. 3rd edn. (reprint of the original 1983) Frankfurter Fachbuchhandlung für Psychologie, Frankfurt
Rosenthal R (1965) Clever Hans: the horse of Mr. Von Osten. Holt, Rinehart and Winston, New York
Uller C, Lewis J (2009) Horses (Equus caballus) select the greater of two quantities in small numerical sets. Anim Cogn 12:733–738 |
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Corporate Author |
Gabor, V. |
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Publisher |
Xenophon Publishing |
Place of Publication |
Wald |
Editor |
; Krueger, K. |
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978-3-95625-000-2 |
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Equine Behaviour @ team @ |
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5894 |
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Author |
Bouskila, A.,; Lourie,E.; de Vries, H.; Hermans, Z.M .; van Dierendonck, M. |
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Title |
Sex, but not relatedness nor age, affect the social network of horses in a semi-natural reserve |
Type |
Conference Article |
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Year |
2015 |
Publication |
Proceedings of the 3. International Equine Science Meeting |
Abbreviated Journal |
Proc. 3. Int. Equine. Sci. Mtg |
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For some time now, social network analysis provides tools to describe concisely the social structure of animals. Nevertheless, the factors that shape the social network and determine the frequency of different types of affiliations are often unclear. For example, the effects of relatedness on social networks have been only studied on relatively few species. Both among social or sub-social species the effects ranged from no effect to a substantial effect. Here we test the hypothesis that relatedness has an effect on the social network of horses (Equus caballus) that live freely in semi-natural conditions, and specifically, that individuals with high values of relatedness to their neighbors in the network will have fewer links Wolf et al, 2011). This hypothesis is based on the notion that related individuals have common interests and are likely to assist each other, and may need less links to other, unrelated individuals. In addition, individuals of similar age and sex are likely to have common needs, and thus are more likely to be associated. We thus tested a second hypothesis: homophily to individuals with similar age and sex will have a role in determining the associations within the social network. The field study was conducted on 27 horses in the Blauwe Kamer reserve in the Netherlands (1.1 sq km). We videotaped horses and their groups and used the information from 22 horses, after excluding the five foals from the analysis. Relatedness was calculated from the pedigree, which was based on parentage, determined by DNA analysis. The social network was constructed based on spatial proximity data. We assessed the influence of relatedness, age-homophily and sex-homophily on the network structure with Multiple Regression Quadratic Assignment Procedure (MRQAP) (Krackhardt 1988), with the R package sna. The results show that there was no significant effect of relatedness on the network, nor an effect of age-homophily. Nevertheless, we found a significant effect of sex-homophily, the tendency of individuals to associate with individuals of the same sex. We argue that the lack of a relatedness effect is not likely to have been caused due to the inability to detect who is kin. The structured social system in horses includes strong associations between often unrelated individuals, alongside with young individuals leaving their natal harem as part of the inherent inbreeding avoidance (Linklater & Cameron 2009; Boyd et al. in press). The significant effect of sex-homophily could stem from the protection females get from associations with other females, e.g., in the reduction of harassment from males. Previous studies on feral horses showed that mares that were better connected with other females in their harem benefited from higher survival rates to their foals (Cameron et al. 2009). The associations among bachelor males could also contribute to the strength of the sex-homophily effect.
In order to generalize from our results, one needs to examine additional populations of horses, because the conditions in the Blauwe Kamer reserve may not be representative, mainly due to the limited opportunity for dispersal in a restricted area.
Keywords: Long-term affiliation; spatial proximity, kin detection
Boyd, L., Scorolli, A. L., Nowzari, H., & Bouskila, A. (2016). Chapter 2: Social organization. In J. I. Ransom, & P. Kaczensky (Eds.), Wild equids. Baltimore, Maryland (in press): The Johns Hopkins University Press.
Cameron, E. Z., Setsaas, T. H., & Linklater, W. L. (2009). Social bonds between unrelated females increase reproductive success in feral horses. Proceedings of the National Academy of Sciences, 106(33), 13850-13853
Krackhardt, D. (1988). Predicting with networks: Nonparametric multiple regression analysis of dyadic data. Social Networks, 10(4), 359-381.
Linklater, W. L., & Cameron, E. Z. (2009). Social dispersal but with philopatry reveals incest avoidance in a polygynous ungulate. Animal Behaviour, 77(5), 1085-1093.
Wolf, J. B., Traulsen, A., & James, R. (2011). Exploring the link between genetic relatedness r and social contact structure k in animal social networks. The American Naturalist, 177(1), 135-142. |
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Bouskila, A. |
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Equine Behaviour @ team @ |
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5881 |
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Siniscalchi, M.; Padalino, B.; Aubé, L.; Quaranta, A. |
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Title |
Right-nostril use during sniffing at arousing stimuli produces higher cardiac activity in jumper horses |
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Journal Article |
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2015 |
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Laterality: Asymmetries of Body, Brain and Cognition |
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Laterality |
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20 |
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4 |
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483-500 |
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Lateralization in horses, Equus caballus, has been reported at both motor and sensory levels. Here we investigated left- and right-nostril use in 12 jumper horses freely sniffing different emotive stimuli. Results revealed that during sniffing at adrenaline and oestrus mare urine stimuli, horses showed a clear right-nostril bias while just a tendency in the use of the right nostril was observed during sniffing of other odours (food, cotton swab and repellent). Sniffing at adrenaline and urine odours was also accompanied by increasing cardiac activity and behavioural reactivity strengthening the role of the right hemisphere in the analysis of intense emotion and sexual behaviour. |
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Routledge |
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1357-650x |
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doi: 10.1080/1357650X.2015.1005629 |
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Equine Behaviour @ team @ |
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6208 |
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Burton, A.C.; Neilson, E.; Moreira, D.; Ladle, A.; Steenweg, R.; Fisher, J.T.; Bayne, E.; Boutin, S. |
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REVIEW: Wildlife camera trapping: a review and recommendations for linking surveys to ecological processes |
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Journal Article |
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2015 |
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Journal of Applied Ecology |
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J Appl Ecol |
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52 |
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3 |
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675-685 |
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animal movement; camera trap; capture-recapture; density estimation; imperfect detection; mammal monitoring; occupancy model; relative abundance; sampling error; wildlife survey methodology |
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Summary Reliable assessment of animal populations is a long-standing challenge in wildlife ecology. Technological advances have led to widespread adoption of camera traps (CTs) to survey wildlife distribution, abundance and behaviour. As for any wildlife survey method, camera trapping must contend with sources of sampling error such as imperfect detection. Early applications focused on density estimation of naturally marked species, but there is growing interest in broad-scale CT surveys of unmarked populations and communities. Nevertheless, inferences based on detection indices are controversial, and the suitability of alternatives such as occupancy estimation is debatable. We reviewed 266 CT studies published between 2008 and 2013. We recorded study objectives and methodologies, evaluating the consistency of CT protocols and sampling designs, the extent to which CT surveys considered sampling error, and the linkages between analytical assumptions and species ecology. Nearly two-thirds of studies surveyed more than one species, and a majority used response variables that ignored imperfect detection (e.g. presence?absence, relative abundance). Many studies used opportunistic sampling and did not explicitly report details of sampling design and camera deployment that could affect conclusions. Most studies estimating density used capture?recapture methods on marked species, with spatially explicit methods becoming more prominent. Few studies estimated density for unmarked species, focusing instead on occupancy modelling or measures of relative abundance. While occupancy studies estimated detectability, most did not explicitly define key components of the modelling framework (e.g. a site) or discuss potential violations of model assumptions (e.g. site closure). Studies using relative abundance relied on assumptions of equal detectability, and most did not explicitly define expected relationships between measured responses and underlying ecological processes (e.g. animal abundance and movement). Synthesis and applications. The rapid adoption of camera traps represents an exciting transition in wildlife survey methodology. We remain optimistic about the technology's promise, but call for more explicit consideration of underlying processes of animal abundance, movement and detection by cameras, including more thorough reporting of methodological details and assumptions. Such transparency will facilitate efforts to evaluate and improve the reliability of camera trap surveys, ultimately leading to stronger inferences and helping to meet modern needs for effective ecological inquiry and biodiversity monitoring. |
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John Wiley & Sons, Ltd |
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0021-8901 |
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https://doi.org/10.1111/1365-2664.12432 |
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Equine Behaviour @ team @ |
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6703 |
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Malavasi, R.; Huber, L. |
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Referential communication in the domestic horse (Equus caballus): first exploration in an ungulate species |
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Conference Article |
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2015 |
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Proceedings of the 3. International Equine Science Meeting |
Abbreviated Journal |
Proc. 3. Int. Equine. Sci. Mtg |
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domestic horse, referential communication, human-horse communication, intentionality |
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An important question in the study of animal communication is whether non-human animals are able to produce communicative gestures, i.e. nonvocal bodily actions directed to a recipient, physically ineffective but with a meaning shared in the social group [1]. Passive gestures are instrumental, tuned to the mere presence/absence of others, whereas active informers recognize receivers as communicative agents and activate shared-attention mechanisms for identifying their attentional state (SAM [2]; e.g. Schwab and Huber [3]). Six operational criteria must be evaluated to classify a signal as referential and intentional [4]: (1) alternative gazes between the partner and the target; (2) apparent attention-getting behaviours are deployed; (3) an audience is required to exhibit the behaviour; (4) the attentional status of an observer influences the propensity to exhibit behaviours; (5) communication is persistent and (6) there is elaboration of communicative behaviour when apparent attempts to manipulate the partner fail. Dogs [5] and non-human primates (reviewed in Liebal and Call [6]) can tune a human receiver’s attention to the object of interest by combining directional and attention-getting signals, such as turning the head or body, gazing to the receiver, and/or establishing eye contact. Research on other species is scarce.
Horses rely on humans to survive in domestic settings and may have evolved skills for communicating flexibly with them [7]. Horses understand human attentional cues (such as body and head orientation, eyes opened/closed) [8], permanent pointing [9] and, to some extent, gazing [10]. Here we tested the ability of 14 outdoor, herd-living domestic horses to communicate referentially with a human partner about the location of a desired target, a bucket of food out of reach. After the baiting of two buckets placed in opposite, unreachable locations were shown by the experimenter, the subject would walk to one of the two buckets. Because approaching a bucket would reveal that the food is out of reach, we expected the horse to look back to the experimenter, then to the bucket, and alternate this gazing several times to indicate its intention. To test whether our prediction is correct and alternate gazing is indeed the result of the horse's referential communication, we video-recorded the behaviour of the subjects in the test (FORWARD) and three control conditions: (1) FORWARD: experimenter oriented to the center of the arena, (2) BACK: experimenter backward oriented in respect to the arena, (3) ALONE: experimenter absent, (4) MANY: as FORWARD plus a familiar human oriented to the subject behind the bucket (Figure 1). We used a conservative criterion of back gazing by considering only turning the head back more than 90 degrees. The results confirmed our prediction. The horses alternated gazes between the partner and the buck significantly more often in the FORWARD than in all the other conditions (Table 1), thus satisfying operational criteria #1, #3 and #4. They also alternated head nods with gazes to the partner significantly more often during the FORWARD condition. We thus considered head nods not an instrumental signal of arousal, but an attention-getting behaviour with communicative function. Subjects used both head nods and neck stretched toward the buck more often in the FORWARD than in the BACK and the ALONE conditions, thus satisfying criteria #2, #3 and #4. In condition MANY, the frequency of head nods did not differ from condition FORWARD, probably because nods were directed to the additional partner behind the buck. This also satisfies criteria #4. The horses gazed to the partner most often in the FORWARD than in the BACK and the MANY conditions, but not in the ALONE. In this condition, subjects could observe the partner walking further from the test arena. To test for the different functions of gazes in presence and in absence of the partner, we compared their average duration between the two conditions: the significantly longer duration of gazes when the subject was alone suggests the instrumental monitoring function of gazes in this experimental condition.
Altogether, the findings suggest that domestic horses possess the ability to use referential communication in an interspecific context, but additional analyses are needed to test for operational criteria #5 and #6. Flexible and voluntary use of communicative signals reveal sophisticated cognitive processes involved in the strategic emission of these signals, and the finding of referential communication skills in an ungulate species forces us to reconsider the evolutionary path of intelligence. Furthermore, ungulates are used intensively by humans (transportation, meat, agriculture, leisure activities), and their welfare is often compromised. Determining whether ungulates can communicate their needs and preferences is paramount to a proper ethical management. |
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Malavasi, R. |
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Equine Behaviour @ team @ |
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5876 |
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Maros, K.; Kovács, R.; Nagy, K. |
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Title |
Questionnaire survey personality assessment of horses of different use |
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2015 |
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Proceedings of the 3. International Equine Science Meeting |
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Proc. 3. Int. Equine. Sci. Mtg |
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horse, personality, questionnaire, |
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We collected data from 248 horses of different breeds, age, sex and use, forming four groups: 74 trotters, 70 gallop horses, 60 horseback archery horses and 44 police horses. All horses were trained and ridden/driven in a regular base. Caretakers or owners who were familiar with the target animals were asked to assess their horses’ temperament. The temperament scores were obtained with the 7-point scale questionnaire according to the Horse Personality Questionnaire which has 25 items and has previously been shown to be reliable for the assessment of personality in horses. It measures six personality components in horses: Dominance, Anxiousness, Excitability, Protection, Sociability and Inquisitiveness.
Component scores were calculated according to Lloyd, A.S., Martin, J.E., Bornett-Gauci, H.L.I., Wilkinson, R.G. (2008) Horse personality: Variation between breeds. Applied Animal Behaviour Science 112. 369–383. The component scores were compared across the four examined groups using the Kruskal–Wallis test. Post hoc multiple comparisons tests were then carried out to explore specific breed differences on each component. The value of alpha was set at 0.05 for all statistical tests.
Groups differed significantly regarding Anxiousness and Excitability, but no significant differences were found regarding Dominance, Protection, Sociability or Inquisitiveness among groups. This finding is in line with the findings of Lloyd et al. (2008) who showed that Anxiousness and Excitability components have the highest level of variation between breeds.In our study, gallop horses had the highest rank regarding Excitability and they differed significantly from police horses which had the lowest rank for this personality component. Interestingly gallop horses had the lowest rank regarding Anxiousness, and trotters got the highest rank in in this component.
According to our results gallop horses are the most extreme in their personality. It is conceivable that being excitable is a more favourable trait for a race horse than for a working police horse. However, it is interesting that trotters are more anxious than gallop horses since they also have a high thoroughbred ancestry. The effect of work and training on these horses needs further surveys. |
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Maros, K. |
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Xenophon Publishing |
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Wald |
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Krueger, K. |
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978-3-95625-000-2 |
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Equine Behaviour @ team @ |
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5904 |
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