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	Author	Proops, L.; McComb, K.
	Title	Cross-modal individual recognition in domestic horses (Equus caballus) extends to familiar humans			Type	Journal Article
	Year	2012	Publication	Proceedings of the Royal Society B: Biological Sciences	Abbreviated Journal
	Volume	279	Issue	1741	Pages	3131-3138
	Keywords
	Abstract	It has recently been shown that some non-human animals can cross-modally recognize members of their own taxon. What is unclear is just how plastic this recognition system can be. In this study, we investigate whether an animal, the domestic horse, is capable of spontaneous cross-modal recognition of individuals from a morphologically very different species. We also provide the first insights into how cross-modal identity information is processed by examining whether there are hemispheric biases in this important social skill. In our preferential looking paradigm, subjects were presented with two people and playbacks of their voices to determine whether they were able to match the voice with the person. When presented with familiar handlers subjects could match the specific familiar person with the correct familiar voice. Horses were significantly better at performing the matching task when the congruent person was standing on their right, indicating marked hemispheric specialization (left hemisphere bias) in this ability. These results are the first to demonstrate that cross-modal recognition in animals can extend to individuals from phylogenetically very distant species. They also indicate that processes governed by the left hemisphere are central to the cross-modal matching of visual and auditory information from familiar individuals in a naturalistic setting.
	Address
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	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
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	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5616
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	Author	McComb, K.; Shannon, G.; Durant, S.M.; Sayialel, K.; Slotow, R.; Poole, J.; Moss, C.
	Title	Leadership in elephants: the adaptive value of age			Type	Journal Article
	Year	2011	Publication	Proceedings of the Royal Society B: Biological Sciences	Abbreviated Journal	Proc. R. Soc. Lond. B.
	Volume	278	Issue	1722	Pages	3270-3276
	Keywords
	Abstract	The value of age is well recognized in human societies, where older individuals often emerge as leaders in tasks requiring specialized knowledge, but what part do such individuals play in other social species? Despite growing interest in how effective leadership might be achieved in animal social systems, the specific role that older leaders may play in decision-making has rarely been experimentally investigated. Here, we use a novel playback paradigm to demonstrate that in African elephants (Loxodonta africana), age affects the ability of matriarchs to make ecologically relevant decisions in a domain critical to survival—the assessment of predatory threat. While groups consistently adjust their defensive behaviour to the greater threat of three roaring lions versus one, families with younger matriarchs typically under-react to roars from male lions despite the severe danger they represent. Sensitivity to this key threat increases with matriarch age and is greatest for the oldest matriarchs, who are likely to have accumulated the most experience. Our study provides the first empirical evidence that individuals within a social group may derive significant benefits from the influence of an older leader because of their enhanced ability to make crucial decisions about predatory threat, generating important insights into selection for longevity in cognitively advanced social mammals.
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	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5652
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	Author	McComb, K.; Clutton-Brock, T.
	Title	Is mate choice copying or aggregation responsible for skewed distributions of females on leks?			Type	Journal Article
	Year	1994	Publication	Proceedings. Biological sciences / The Royal Society	Abbreviated Journal	Proc Biol Sci
	Volume	255	Issue	1342	Pages	13-19
	Keywords	Animals; Deer/physiology; Estrus/physiology; Female; Male; Phenotype; Sexual Behavior, Animal/physiology; Territoriality
	Abstract	In several lek-breeding populations of birds and mammals, females arriving on leks tend to join males that already have females in their territories. This might occur either because females have an evolved preference for mating with males that are attractive to other females, or because they join groups of other females to obtain greater safety from predation or dangerous harassment by males. We have previously used controlled experiments to show that oestrous fallow deer females join males with established harems because they are attracted to female groups rather than to the males themselves. Here we demonstrate that the preference for males with females over males without females is specific to oestrous females and weak or absent in anoestrous ones, and that it is not associated with a preference for mating with males that have previously been seen to mate with other females. Furthermore, oestrous females given the choice between males that do not already have females with them show no significant preference for antlered over deantlered males or for older males over younger ones. We conclude that female attraction to other females on the lek is likely to be an adaptation to avoiding harassment in mixed-sex herds. In this situation, a male's ability to maintain the cohesion of his harem may be the principal cause of variation in mating success between males.
	Address	Department of Zoology, University of Cambridge, U.K
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0962-8452	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:8153135			Approved	no
	Call Number				Serial	1823
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	Author	Proops, L.; McComb, K.; Reby, D.
	Title	*Cross-modal individual recognition in domestic horses (Equus caballus)*			Type	Journal Article
	Year	2009	Publication	Proceedings of the National Academy of Sciences of the United States of America	Abbreviated Journal	Proc. Natl. Acad. Sci. U.S.A.
	Volume	106	Issue	3	Pages	947-951
	Keywords	animal cognition vocal communication social behavior playback experiment expectancy violation
	Abstract	Individual recognition is considered a complex process and, although it is believed to be widespread across animal taxa, the cognitive mechanisms underlying this ability are poorly understood. An essential feature of individual recognition in humans is that it is cross-modal, allowing the matching of current sensory cues to identity with stored information about that specific individual from other modalities. Here, we use a cross-modal expectancy violation paradigm to provide a clear and systematic demonstration of cross-modal individual recognition in a nonhuman animal: the domestic horse. Subjects watched a herd member being led past them before the individual went of view, and a call from that or a different associate was played from a loudspeaker positioned close to the point of disappearance. When horses were shown one associate and then the call of a different associate was played, they responded more quickly and looked significantly longer in the direction of the call than when the call matched the herd member just seen, an indication that the incongruent combination violated their expectations. Thus, horses appear to possess a cross-modal representation of known individuals containing unique auditory and visual/olfactory information. Our paradigm could provide a powerful way to study individual recognition across a wide range of species.
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	Notes	10.1073/pnas.0809127105			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	4689
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	Author	Proops, L.; Walton, M.; McComb, K.
	Title	The use of human-given cues by domestic horses, Equus caballus, during an object choice task			Type	Journal Article
	Year	2010	Publication	Animal Behaviour.	Abbreviated Journal	Anim. Behav.
	Volume	79	Issue	6	Pages	1205-1209
	Keywords	domestication; Equus caballus; horse; interspecific communication; marker cue; pointing; social cognition
	Abstract	Selection pressures during domestication are thought to lead to an enhanced ability to use human-given cues. Horses fulfil a wide variety of roles for humans and have been domesticated for at least 5000 years but their ability to read human cues has not been widely studied. We tested the ability of 28 horses to attend to human-given cues in an object choice task. We included five different cues: distal sustained pointing, momentary tapping, marker placement, body orientation and gaze (head) alternation. Horses were able to use the pointing and marker placement cues spontaneously but not the tapping, body orientation and gaze alternation cues. The overall pattern of responding suggests that horses may use cues that provide stimulus enhancement at the time of choice and do not have an understanding of the communicative nature of the cues given. As such, their proficiency at this task appears to be inferior to that of domestic dogs, Canis lupus familiaris, but similar to that of domestic goats, Caprus hircus.
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	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
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	Series Volume		Series Issue		Edition
	ISSN	0003-3472	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5083
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	Author	McComb, K.; Moss, C.; Sayialel, S.; Baker, L.
	Title	Unusually extensive networks of vocal recognition in African elephants			Type	Journal Article
	Year	2000	Publication	Anim Behav	Abbreviated Journal
	Volume	59	Issue		Pages
	Keywords
	Abstract
	Address
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	Notes				Approved	no
	Call Number	Equine Behaviour @ team @ McComb2000			Serial	6281
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	Author	Proops, L.; Grounds, K.; Smith, A.V.; McComb, K.
	Title	Animals Remember Previous Facial Expressions that Specific Humans Have Exhibited			Type	Journal Article
	Year	2018	Publication	Current Biology	Abbreviated Journal
	Volume	28	Issue	9	Pages	1428-1432.e4
	Keywords	affective processing; face processing; ; animal-human interaction; interspecific communication; animal memory
	Abstract	Summary For humans, facial expressions are important social signals, and how we perceive specific individuals may be influenced by subtle emotional cues that they have given us in past encounters. A wide range of animal species are also capable of discriminating the emotions of others through facial expressions [1, 2, 3, 4, 5], and it is clear that remembering emotional experiences with specific individuals could have clear benefits for social bonding and aggression avoidance when these individuals are encountered again. Although there is evidence that non-human animals are capable of remembering the identity of individuals who have directly harmed them [6, 7], it is not known whether animals can form lasting memories of specific individuals simply by observing subtle emotional expressions that they exhibit on their faces. Here we conducted controlled experiments in which domestic horses were presented with a photograph of an angry or happy human face and several hours later saw the person who had given the expression in a neutral state. Short-term exposure to the facial expression was enough to generate clear differences in subsequent responses to that individual (but not to a different mismatched person), consistent with the past angry expression having been perceived negatively and the happy expression positively. Both humans were blind to the photograph that the horses had seen. Our results provide clear evidence that some non-human animals can effectively eavesdrop on the emotional state cues that humans reveal on a moment-to-moment basis, using their memory of these to guide future interactions with particular individuals.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0960-9822	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6394
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	Author	Wathan, J.; McComb, K.
	Title	The eyes and ears are visual indicators of attention in domestic horses			Type	Journal Article
	Year	2014	Publication	Current Biology	Abbreviated Journal
	Volume	24	Issue	15	Pages	R677-R679
	Keywords
	Abstract	Summary Sensitivity to the attentional states of others has adaptive advantages [1], and in social animals, attending to others is important for predator detection, as well as a pre-requisite for normal social functioning and more complex socio-cognitive abilities [2]. Despite widespread interest in how social species perceive attention in others, studies of non-human animals have been inconclusive about the detailed cues involved [3]. Previous work has focused on head and eye direction, overlooking the fact that many mammals have obvious and mobile ears that could act as a visual cue to attention. Here we report that horses use the head orientation of a conspecific to locate food, but that this ability is disrupted when parts of the face (the eyes and ears) are covered up with naturalistic masks. The ability to correctly judge attention also interacted with the identity of the model horse, suggesting that individual differences in facial features may influence the salience of cues. Our results indicate that a combination of head orientation with facial expression, specifically involving both the eyes and ears, is necessary for communicating social attention. These findings emphasise that in order to understand how attention is communicated in non-human animals, it is essential to consider a broad range of cues.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0960-9822	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5929
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	Author	Proops, L.; McComb, K.
	Title	*Attributing attention: the use of human-given cues by domestic horses (Equus caballus)*			Type	Journal Article
	Year	2010	Publication	Animal Cognition	Abbreviated Journal	Anim. Cogn.
	Volume	13	Issue	2	Pages	197-205
	Keywords
	Abstract	Abstract Recent research has shown that domestic dogs are particularly good at determining the focus of human attention, often outperforming chimpanzees and hand-reared wolves. It has been suggested that the close evolutionary relationship between humans and dogs has led to the development of this ability; however, very few other domestic species have been studied. We tested the ability of 36 domestic horses to discriminate between an attentive and inattentive person in determining whom to approach for food. The cues provided were body orientation, head orientation or whether the experimenters’ eyes were open or closed. A fourth, mixed condition was included where the attentive person stood with their body facing away from the subjects but their head turned towards the subject while the inattentive person stood with their body facing the subject but their head turned away. Horses chose the attentive person significantly more often using the body cue, head cue, and eye cue but not the mixed cue. This result suggests that domestic horses are highly sensitive to human attentional cues, including gaze. The possible role of evolutionary and environmental factors in the development of this ability is discussed.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	4984
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	Author	Smith, A.V.; Proops, L.; Grounds, K.; Wathan, J.; McComb, K.
	Title	Functionally relevant responses to human facial expressions of emotion in the domestic horse (Equus caballus)			Type	Journal Article
	Year	2016	Publication	Biology Letters	Abbreviated Journal	Biol. Lett.
	Volume	12	Issue	2	Pages
	Keywords
	Abstract	Whether non-human animals can recognize human signals, including emotions, has both scientific and applied importance, and is particularly relevant for domesticated species. This study presents the first evidence of horses' abilities to spontaneously discriminate between positive (happy) and negative (angry) human facial expressions in photographs. Our results showed that the angry faces induced responses indicative of a functional understanding of the stimuli: horses displayed a left-gaze bias (a lateralization generally associated with stimuli perceived as negative) and a quicker increase in heart rate (HR) towards these photographs. Such lateralized responses towards human emotion have previously only been documented in dogs, and effects of facial expressions on HR have not been shown in any heterospecific studies. Alongside the insights that these findings provide into interspecific communication, they raise interesting questions about the generality and adaptiveness of emotional expression and perception across species.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6010
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