Monamie Ringhofer
Current Organizations / Institutes
The organizations/institutes from the scholar's latest article
Research Subjects
Monamie Ringhofer has a total of 32 co-authors. Here are the top co-authors by number of shared publications:
Shinya Yamamoto
Institute for Advanced Study, Kyoto University, Kyoto, Japan
5 shared publications- Shinya Yamamoto
Kyoto University
5 shared publications Sota Inoue
Wildlife Research Center, Kyoto University, Kyoto, Japan
4 shared publicationsKazushi Ikeda
Nara Institute of Science and Technology 2
3 shared publications- Renata S. Mendonça
Primate Research Institute, Kyoto University, Inuyama, Aichi, 484-8506, Japan.
3 shared publications Satoshi Hirata
Wildlife Research Center, Kyoto University, Kyoto, Japan
3 shared publications
Latest Publications
- Open Access
- 21 January 2026
- Published by The Royal Society in Proceedings Of The Royal Society B-Biological Sciences
- Vol. 293 (2063)
Spatial strategies in non-territorial societies: how feral horses maintain boundaries with other groups
Encounters between competitive individuals or groups are common in social animals and can involve costly aggression; thus, animals often employ strategies to minimize direct conflict. However, research on whether and how animals adjust their group spatial structure when they encounter or spatially co-occur with a different group remains limited. We investigated how non-territorial units in feral horse multilevel societies manage spatial encounters with neighbouring units. We observed 25 reproductive units in northern Portugal, using drones, and employed statistical analyses to quantify spatial dynamics. We found that horses actively adjusted their spatial formations depending on the proximity of other units. Units became more circular and cohesive when other units approached, but when extremely close, they elongated their shapes to avoid boundary crossings. These adjustments indicate that horses maintain flexible unit boundaries to prevent mixing with other units. A notable exception was a particular pair of units that frequently crossed boundaries and intermixed, representing a unique social level within the horse multilevel society. Overall, our findings indicate that feral horses respond adaptively to inter-unit encounters by balancing the benefits of aggregation with the need to reduce direct overlap, providing new insights into the spatial organization of non-territorial societies.
- 21 January 2026
- Published by The Royal Society in Proceedings Of The Royal Society B-Biological Sciences
- Vol. 293 (2063)
Open AccessSpatial strategies in non-territorial societies: how feral horses maintain boundaries with other groups
Encounters between competitive individuals or groups are common in social animals and can involve costly aggression; thus, animals often employ strategies to minimize direct conflict. However, research on whether and how animals adjust their group spatial structure when they encounter or spatially co-occur with a different group remains limited. We investigated how non-territorial units in feral horse multilevel societies manage spatial encounters with neighbouring units. We observed 25 reproductive units in northern Portugal, using drones, and employed statistical analyses to quantify spatial dynamics. We found that horses actively adjusted their spatial formations depending on the proximity of other units. Units became more circular and cohesive when other units approached, but when extremely close, they elongated their shapes to avoid boundary crossings. These adjustments indicate that horses maintain flexible unit boundaries to prevent mixing with other units. A notable exception was a particular pair of units that frequently crossed boundaries and intermixed, representing a unique social level within the horse multilevel society. Overall, our findings indicate that feral horses respond adaptively to inter-unit encounters by balancing the benefits of aggregation with the need to reduce direct overlap, providing new insights into the spatial organization of non-territorial societies.
- 17 December 2025
- Published by The Royal Society in Journal of The Royal Society Interface
- Vol. 22 (233)
Herding as an emergent behaviour in harem groups of feral Garrano ponies
Collective decision-making and movement coordination are essential behaviours observed in biological systems, from animal herds to human crowds. Horses are a highly social species with a multilevel society. Herding, where the harem is collected to move in a certain direction, is an often-cited example of agonistic behaviour in horses, yet poorly understood in a granular, quantitative sense. We use transfer entropy to measure herding in a harem group of feral Garrano ponies in Serra D’Arga, Portugal. First, we characterize the harem’s leader–follower relationships by quantifying the time lag (average 1.44 s) and duration (average 1.72 s) of influence during herding, establishing variance across social characteristics. Second, we internally validate transfer entropy as a method to detect herding by comparing it with traditional clustering methods. To augment the paucity of existing data, synthetic data is generated from a mathematical model of feral horse harems, demonstrating superior accuracy (0.80) and F1-score (0.76) against traditional clustering and time-series synchrony methods. Third, we provide evidence for herding as an emergent behaviour: leadership influence often propagates indirectly among mares in short bursts of information flow before reaching the entire harem. These results enrich our understanding of horse behaviour and provide a foundation for using transfer entropy to study decision-making in other species.
- 17 December 2025
- Published by The Royal Society in Journal of The Royal Society Interface
- Vol. 22 (233)
Herding as an emergent behaviour in harem groups of feral Garrano ponies
Collective decision-making and movement coordination are essential behaviours observed in biological systems, from animal herds to human crowds. Horses are a highly social species with a multilevel society. Herding, where the harem is collected to move in a certain direction, is an often-cited example of agonistic behaviour in horses, yet poorly understood in a granular, quantitative sense. We use transfer entropy to measure herding in a harem group of feral Garrano ponies in Serra D’Arga, Portugal. First, we characterize the harem’s leader–follower relationships by quantifying the time lag (average 1.44 s) and duration (average 1.72 s) of influence during herding, establishing variance across social characteristics. Second, we internally validate transfer entropy as a method to detect herding by comparing it with traditional clustering methods. To augment the paucity of existing data, synthetic data is generated from a mathematical model of feral horse harems, demonstrating superior accuracy (0.80) and F1-score (0.76) against traditional clustering and time-series synchrony methods. Third, we provide evidence for herding as an emergent behaviour: leadership influence often propagates indirectly among mares in short bursts of information flow before reaching the entire harem. These results enrich our understanding of horse behaviour and provide a foundation for using transfer entropy to study decision-making in other species.
- 18 March 2025
- Published by Elsevier in Animal Behaviour
- Vol. 222
No evidence of kin preferences in a feral horse multilevel society
- 18 March 2025
- Published by Elsevier in Animal Behaviour
- Vol. 222
No evidence of kin preferences in a feral horse multilevel society
- 13 August 2022
- Published by Springer Nature in Animal Cognition
- Vol. 26 (2) , 369-377
Horses form cross-modal representations of adults and children
Recently, research on domestic mammals’ sociocognitive skills toward humans has been prolific, allowing us to better understand the human–animal relationship. For example, horses have been shown to distinguish human beings on the basis of photographs and voices and to have cross-modal mental representations of individual humans and human emotions. This leads to questions such as the extent to which horses can differentiate human attributes such as age. Here, we tested whether horses discriminate human adults from children. In a cross-modal paradigm, we presented 31 female horses with two simultaneous muted videos of a child and an adult saying the same neutral sentence, accompanied by the sound of an adult’s or child’s voice speaking the sentence. The horses looked significantly longer at the videos that were incongruent with the heard voice than at the congruent videos. We conclude that horses can match adults’ and children’s faces and voices cross-modally. Moreover, their heart rates increased during children’s vocalizations but not during adults’. This suggests that in addition to having mental representations of adults and children, horses have a stronger emotional response to children’s voices than adults’ voices.
- 13 August 2022
- Published by Springer Nature in Animal Cognition
- Vol. 26 (2) , 369-377
Horses form cross-modal representations of adults and children
Recently, research on domestic mammals’ sociocognitive skills toward humans has been prolific, allowing us to better understand the human–animal relationship. For example, horses have been shown to distinguish human beings on the basis of photographs and voices and to have cross-modal mental representations of individual humans and human emotions. This leads to questions such as the extent to which horses can differentiate human attributes such as age. Here, we tested whether horses discriminate human adults from children. In a cross-modal paradigm, we presented 31 female horses with two simultaneous muted videos of a child and an adult saying the same neutral sentence, accompanied by the sound of an adult’s or child’s voice speaking the sentence. The horses looked significantly longer at the videos that were incongruent with the heard voice than at the congruent videos. We conclude that horses can match adults’ and children’s faces and voices cross-modally. Moreover, their heart rates increased during children’s vocalizations but not during adults’. This suggests that in addition to having mental representations of adults and children, horses have a stronger emotional response to children’s voices than adults’ voices.
- Open Access
- 10 August 2021
- Published by Springer Nature in Scientific Reports
- Vol. 11 (1) , 1-9
Horses with sustained attention follow the pointing of a human who knows where food is hidden
When interacting with humans, domesticated species may respond to communicative gestures, such as pointing. However, it is currently unknown, except for in dogs, if species comprehend the communicative nature of such cues. Here, we investigated whether horses could follow the pointing of a human informant by evaluating the credibility of the information about the food-hiding place provided by the pointing of two informants. Using an object-choice task, we manipulated the attentional state of the two informants during food-hiding events and differentiated their knowledge about the location of the hidden food. Furthermore, we investigated the horses’ visual attention levels towards human behaviour to evaluate the relationship between their motivation and their performance of the task. The result showed that horses that sustained high attention levels could evaluate the credibility of the information and followed the pointing of an informant who knew where food was hidden (Z = − 2.281, P = 0.002, n = 36). This suggests that horses are highly sensitive to the attentional state and pointing gestures of humans, and that they perceive pointing as a communicative cue. This study also indicates that the motivation for the task should be investigated to determine the socio-cognitive abilities of animals.
- 10 August 2021
- Published by Springer Nature in Scientific Reports
- Vol. 11 (1) , 1-9
Open AccessHorses with sustained attention follow the pointing of a human who knows where food is hidden
When interacting with humans, domesticated species may respond to communicative gestures, such as pointing. However, it is currently unknown, except for in dogs, if species comprehend the communicative nature of such cues. Here, we investigated whether horses could follow the pointing of a human informant by evaluating the credibility of the information about the food-hiding place provided by the pointing of two informants. Using an object-choice task, we manipulated the attentional state of the two informants during food-hiding events and differentiated their knowledge about the location of the hidden food. Furthermore, we investigated the horses’ visual attention levels towards human behaviour to evaluate the relationship between their motivation and their performance of the task. The result showed that horses that sustained high attention levels could evaluate the credibility of the information and followed the pointing of an informant who knew where food was hidden (Z = − 2.281, P = 0.002, n = 36). This suggests that horses are highly sensitive to the attentional state and pointing gestures of humans, and that they perceive pointing as a communicative cue. This study also indicates that the motivation for the task should be investigated to determine the socio-cognitive abilities of animals.
- Open Access
- 5 January 2021
- Published by Springer Nature in Journal of Ethology
- Vol. 39 (1) , 145
Correction to: Herding mechanisms to maintain the cohesion of a harem group: two interaction phases during herding
The article Herding mechanisms to maintain the cohesion of a harem group.
- 5 January 2021
- Published by Springer Nature in Journal of Ethology
- Vol. 39 (1) , 145
Open AccessCorrection to: Herding mechanisms to maintain the cohesion of a harem group: two interaction phases during herding
The article Herding mechanisms to maintain the cohesion of a harem group.
- Open Access
- 11 November 2019
- Published by Springer Nature in Journal of Ethology
- Vol. 38 (1) , 71-77
Herding mechanisms to maintain the cohesion of a harem group: two interaction phases during herding
In animal groups, individual interactions achieve coordinated movements to maintain cohesion. In horse harem groups, herding is a behavior in which males chase females from behind; it is considered to assist with group cohesiveness. However, the mechanisms by which the individuals move to maintain group cohesion are unknown. We applied novel non-invasive methods of drone filming and video tracking to observe horse movements in the field with high temporal and spatial resolution. We tracked all group members and drew trajectories. We analyzed the movements of females and found two phases of interactions based on their timing of movement initiation. The females that moved first were those nearest to the herding male, while the movement initiation of the later females was determined by the distance from the nearest moving female, not by the distance from the herding male. These interactions are unique among animal group movements and might represent a herding mechanism responsible for maintaining group cohesion. This might be due to long-term stable relationships within a harem group and strong social bonds between females. This study showed that the combination of drone filming and video tracking is a useful method for analyzing the movements of animals simultaneously in high resolution.
- 11 November 2019
- Published by Springer Nature in Journal of Ethology
- Vol. 38 (1) , 71-77
Open AccessHerding mechanisms to maintain the cohesion of a harem group: two interaction phases during herding
In animal groups, individual interactions achieve coordinated movements to maintain cohesion. In horse harem groups, herding is a behavior in which males chase females from behind; it is considered to assist with group cohesiveness. However, the mechanisms by which the individuals move to maintain group cohesion are unknown. We applied novel non-invasive methods of drone filming and video tracking to observe horse movements in the field with high temporal and spatial resolution. We tracked all group members and drew trajectories. We analyzed the movements of females and found two phases of interactions based on their timing of movement initiation. The females that moved first were those nearest to the herding male, while the movement initiation of the later females was determined by the distance from the nearest moving female, not by the distance from the herding male. These interactions are unique among animal group movements and might represent a herding mechanism responsible for maintaining group cohesion. This might be due to long-term stable relationships within a harem group and strong social bonds between females. This study showed that the combination of drone filming and video tracking is a useful method for analyzing the movements of animals simultaneously in high resolution.
- 1 September 2019
- Published by Elsevier in Behavioural Processes
- Vol. 166, 103906
Horses prefer to solicit a person who previously observed a food-hiding process to access this food: A possible indication of attentional state attribution
- 1 September 2019
- Published by Elsevier in Behavioural Processes
- Vol. 166, 103906
Horses prefer to solicit a person who previously observed a food-hiding process to access this food: A possible indication of attentional state attribution
- 27 May 2019
- Published by Springer Nature in Primates
- Vol. 61 (1) , 49-54
Feral horses’ (Equus ferus caballus) behavior toward dying and dead conspecifics
In the rapidly expanding field of comparative thanatology, reports from a wide range of taxa suggest that some aspects of a concept of death may be shared by many non-human species. In horses, there are only a few anecdotal reports on behaviors toward dead conspecifics, mostly concerning domestic individuals. Here, we describe the case of a 2-month-old, free-ranging male foal that died around 12 h after being found severely injured due to a presumed wolf attack, focusing on other individuals’ reactions to the dying foal. We also placed camera traps near horse carcasses to investigate reactions by other horses. Kin and non-kin of both sexes showed unusual interest in the dying foal. However, horses appeared to avoid dead conspecifics. Recording individual reactions to dead and dying conspecifics in naturalistic settings will enhance our knowledge about death-related behaviors in horses, allowing comparisons with other species that have been more thoroughly studied, to understand the evolutionary basis of these behaviors.
- 27 May 2019
- Published by Springer Nature in Primates
- Vol. 61 (1) , 49-54
Feral horses’ (Equus ferus caballus) behavior toward dying and dead conspecifics
In the rapidly expanding field of comparative thanatology, reports from a wide range of taxa suggest that some aspects of a concept of death may be shared by many non-human species. In horses, there are only a few anecdotal reports on behaviors toward dead conspecifics, mostly concerning domestic individuals. Here, we describe the case of a 2-month-old, free-ranging male foal that died around 12 h after being found severely injured due to a presumed wolf attack, focusing on other individuals’ reactions to the dying foal. We also placed camera traps near horse carcasses to investigate reactions by other horses. Kin and non-kin of both sexes showed unusual interest in the dying foal. However, horses appeared to avoid dead conspecifics. Recording individual reactions to dead and dying conspecifics in naturalistic settings will enhance our knowledge about death-related behaviors in horses, allowing comparisons with other species that have been more thoroughly studied, to understand the evolutionary basis of these behaviors.
- 5 June 2017
- Published by Springer Nature in Primates
- Vol. 58 (4) , 479-484
Comparison of the social systems of primates and feral horses: data from a newly established horse research site on Serra D’Arga, northern Portugal
Horses are phylogenetically distant from primates, but considerable behavioral links exist between the two. The sociality of horses, characterized by group stability, is similar to that of primates, but different from that of many other ungulates. Although horses and primates are good models for exploring the evolution of societies in human and non-human animals, fewer studies have been conducted on the social system of horses than primates. Here, we investigated the social system of feral horses, particularly the determinant factors of single-male/multi-male group dichotomy, in light of hypotheses derived from studies of primate societies. Socioecological data from 26 groups comprising 208 feral horses on Serra D’Arga, northern Portugal suggest that these primate-based hypotheses cannot adequately explain the social system of horses. In view of the sympatric existence of multi- and single-male groups, and the frequent intergroup transfers and promiscuous mating of females with males of different groups, male–female relationships of horses appear to differ from those of polygynous primates.
- 5 June 2017
- Published by Springer Nature in Primates
- Vol. 58 (4) , 479-484
Comparison of the social systems of primates and feral horses: data from a newly established horse research site on Serra D’Arga, northern Portugal
Horses are phylogenetically distant from primates, but considerable behavioral links exist between the two. The sociality of horses, characterized by group stability, is similar to that of primates, but different from that of many other ungulates. Although horses and primates are good models for exploring the evolution of societies in human and non-human animals, fewer studies have been conducted on the social system of horses than primates. Here, we investigated the social system of feral horses, particularly the determinant factors of single-male/multi-male group dichotomy, in light of hypotheses derived from studies of primate societies. Socioecological data from 26 groups comprising 208 feral horses on Serra D’Arga, northern Portugal suggest that these primate-based hypotheses cannot adequately explain the social system of horses. In view of the sympatric existence of multi- and single-male groups, and the frequent intergroup transfers and promiscuous mating of females with males of different groups, male–female relationships of horses appear to differ from those of polygynous primates.