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Novel observation of play behaviour between a harem holder and a bachelor group of Przewalski’s horses in the wild

Novel observation of play behaviour between a harem holder and a bachelor group of Przewalski’s... Przewalski’s horses live in stable nonterritorial families (harem) of one or more harem stallions, several mares, and their offspring. The harem stallion typically behaves aggressively towards any male intruder approaching the harem. Play behav - iour is frequently observed among the group members in horses. For stallions, the most common, well-known cases of play behaviour are found between members of bachelor groups (groups of young stallions) and between the harem stallion and his offspring. The play between the harem stallion and members of the bachelor groups is, on the other hand, seemingly much rarer as a description of such events in the literature is anecdotal. In this note, we present our observation conducted in the Great Gobi B Strictly Protected Area (Mongolia) and describe the play behaviour between a Przewalski’s horse harem stallion and members of a bachelor group. The observation was done as a part of broader research during which selected Przewalski’s horses’ groups were located by binoculars in daily monitoring routines and filmed from a close distance (from 150 to 800 m). Behavioural Observation Research Interactive Software (BORIS) was used to extract data from the video recordings. The observation described in this note lasted for ∼ 180 min, during which the horses engaged in three separate bouts of play and repeated inter-group association. During the whole study (241.5 h of video recordings), this was the only observation including inter-group interactions. We observed two other events during which two harems approached each other (but never merged). Our observation is the first video recording of such an event and raises the question of how preva- lent this behaviour is in the Przewalski’s horses’ restored natural populations. We conclude that this behaviour is unusual or has not been sufficiently studied in wild or feral harems. Further technological advances may help reveal more information on this topic. Keywords Equus ferus przewalskii · Gobi · Stallion · Social behaviour · Mongolia Introduction 1985; Paquette 1994). However, the fact that play is observed in many species after sexual maturity suggests that it may be Since the earliest systematic studies of behaviour, the subject relevant for animals throughout their lifetime (Asensio et al. of why animals play has been a topic of discussion (Darwin 2022; Beckel 1991; Bond and Diamond 2003). 1871; Groos 1911). Play is mainly perceived as a tool by which Adult-only social play is typically observed among group young animals acquire skills (e.g. fighting, mating, social mates and is mainly described as a mechanism for foster- behaviour) necessary for their adult life (Martin and Caro ing social networks and encouraging collaboration among individuals. Adult play may provide a social bonding pur- pose similar to grooming, allowing animals to socialise. In other words, it enables animals to create and sustain social * Francisco Ceacero ceacero@ftz.czu.cz connections as well as restrain anger, lower tension, and enhance tolerance under challenging situations (Baldwin Faculty of Tropical AgriSciences, Czech University of Life and Baldwin 1973; Enomoto 1990; Ferguson and Frankis Sciences Prague, Prague, Czech Republic 2001; Martin and Caro 1985; Merrick 1977; Norscia and Great Gobi B Strictly Protected Area, Takhin Tal, Mongolia Palagi 2011; Palagi et al. 2006). Zoo Praha, U Trojského Zámku 120/3, Prague, Czech Republic Vol.:(0123456789) 1 3 acta ethologica In horses, play is most common and noticeable in foals stallion and members of a bachelor group in the Great Gobi and yearlings, although it also occurs in adult individu- B Strictly Protected Area (Mongolia) to raise the question of als (McDonnell and Poulin 2002). Play is also frequently how prevalent and important this behaviour is in the Przew- observed among the members of the horse bachelor groups alski’s horses’ restored natural populations and to encourage (Feh 2005), including Przewalski’s horses (Christensen et al. future research on this topic. 2002). Play behaviour is believed to be essential in devel- oping specific social skills (e.g. fighting) in young males compared to young females (Crowell-Davis et al. 1987; Rho Methods and results et al. 2007). Indeed, it has been observed that colts (male foals) interact more actively with the harem stallion (their This observation was conducted in July 2018 in the Great father) than fillies (female foals; Crowell-Davis et al. 1987; Gobi B Strictly Protected Area (GGBSPA), Mongolia, dur- Šandlová et al. 2020). However, in adult stallions with their ing broader research fieldwork (July 2018, May–June 2019, harem, observations of play with other mature stallions are and September–October 2019) focusing on the behavioural very scarce (Berger 1986; McDonnell and Poulin 2002). ecology of Przewalski’s horses. In the summer of 2018, two Przewalski’s horses (Equus ferus przewalskii) live in stable, wild-born harems involving 21 adult horses were recorded. nonterritorial family units called harems (Bourjade et al. 2009) In spring 2019, three groups were recorded (two wild-born and exhibit female defence polygyny. The harems consist of harems and one harem consisting of long-term reintroduced one or more harem stallions (males), several mares (females), mares and a wild-born stallion) involving 25 adult horses. and their young progeny (King et al. 2015; Rubenstein 2011). In autumn 2019, four groups were recorded (two wild-born When a colt or filly reaches reproductive age, typically between harems and two harems consisting of long-term reintroduced the ages of one and four, they separate from their family. While mares and wild-born stallions) involving 27 adult horses colts usually join a bachelor group until they can form their (Bernátková et al. 2022). own harem, fillies typically join another stable family unit During the whole study (241.5  h of recordings), we (Bahloul et al. 2001; Boyd and Bandi 2002; International observed only one inter-group interaction (the one described Takhi Group 2021; King et al. 2015). Mature stallions form a in this short note). Once, two harems were observed feeding new harem by taking over an existing harem or joining females close to each other (∼ 50 m) but never merged or interacted dispersed from other harems. These stallions become harem (one of these harems was Tsetsen, the one also involved in holders and protect the mares and their offspring from other the observation described here). Two harems approaching stallions and predators (Boyd et al. 2016). To guard the harem each other (neither of which was Tsetsen) were also observed successfully, the stallion typically stays more vigilant, feeds just once, but both groups stopped ∼ 300 m from each other less, moves more, and shows less behavioural synchrony than except for the harem stallions, who kept approaching each other harem members (King et al. 2016; Souris et al. 2007). other, stopped ∼ 50 m from each other, and returned to their As mares are generally protected from stallion harassment by group. the group structure, they are often in better condition, and their The GGBSPA, in SW Mongolia, is a reintroduction site reproductive success is higher than in mares not bonded to a for Przewalski’s horse and an important refuge for several breeding group (Linklater et al. 1999). It was also hypothesised other endangered species (International Takhi Group 2021; that affiliative interactions between mares and other mares or Kaczensky et al. 2008, 2011). Since 2019, it has encom- stallions might boost reproductive success by minimising har- passed ≈18 000 km of desert steppe and desert habitat assment (Cameron et al. 2009; Linklater et al. 1999). Cases (≈9 000 km before the enlargement, International Takhi of two or more stallions defending the same harem have been Group 2021). At the time of the research, there were 349 reported (Linklater et al. 2013). Previously, it was suggested Przewalski’s horses in 24 harems and 3–5 bachelor groups. that these stallion alliances improve the ability of stallions to The structure and number of the bachelor groups change defend their mutual family group from other stallions (Boyd frequently. During this research, the GGBSPA population et al. 2016) and might increase the foal survival rate in their comprised a maximum of 200 females and 149 males (GGB- group (Feh 1999). However, removal experiments showed that SPA administration, personal communication). Fieldwork the cooperative hypotheses do not appear to explain the exist- consisted of locating Przewalski’s horses’ groups with bin- ence of multi-stallion groups (Linklater et al. 2013). oculars during daily monitoring routines. Once one of the It has been described that in free-roaming populations target groups was localised, it was approached and filmed of horses, most stallions know each other and their rela- from a close distance without disturbing them (from 150 tive fighting ability (Rubenstein and Hack 1992), and most to 800 m) by a 4 K Panasonic VX1 video camera with a encounters result in ritualised behaviours (Sigurjonsdottir tripod. The ethogram consisting of 5 categories (feeding, et al. 2012). In this note, we provide a recording and descrip- locomotion, resting, social, and other) was used to catego- tion of the play behaviour between Przewalski’s horse harem rise observed behaviours. Play behaviour was included in 1 3 acta ethologica the social category (Bernátková et al. 2022). Behavioural 20 m from each other. Eighteen minutes after our observa- Observation Research Interactive Software (BORIS) was tion started, Tsetsen joined the bachelor group again. We used for event logging, coding, and observation of the cap- observed mutual investigation and play behaviour (muzzle tured videos (Friard and Gamba 2016). sniffing, slight rearing, head resting) between him and the Play between harem stallions and members of bachelor stallions of the bachelor group. No direct aggression was groups has never been observed in the population of the observed (video 2). Twenty minutes after our observation GGBSPA (GGBSPA administration, personal commu- started, one 2-year-old stallion from the Tsetsen harem nication). While observing one of the selected groups (a approached the bachelor group and Tsetsen. The rest of wild-born harem called Tsetsen harem), we noticed the non- the harem followed him. Both groups merged momentar- agonistic social behaviour of the harem stallion towards a ily, but after a few mutual aggressive encounters started bachelor group which approached his harem. The Tsetsen by the harem mares (kick threats, kicking), they split. At harem consisted of 8 individuals: 1 harem stallion (Tset- this moment, Tsetsen herded his harem and then chased the sen, 7-year-old), three adult mares (2-, 5-, and 6-year-old), bachelors away. However, 25 min after the start of our obser- 2 2-year-old males, 1 1-year-old male, and 1 1-year-old vation, the Tsetsen harem and bachelor group merged again, female. The GGBSPA administration confirmed that none of and no aggressive behaviour, herding, or any other behaviour the mares was in oestrus during the observation. Tsetsen was aiming to prevent this from the Tsetsen side was observed a harem holder for the first time in his life, and the harem this time (video 3). We observed both groups merging and was relatively new (it was formed just two months before our splitting away multiple times. During these events, Tsetsen observation). All the breeding mares remained there until mostly remained closer to the bachelor group than to his our following fieldwork one year later. Tsetsen established harem. The 2-year-old stallion from the Tsetsen harem was his harem by taking over half of the mares from his father’s also observed joining the bachelor group and Tsetsen in the harem, in which all the mares of our study had been together mutual investigation and play behaviour (video 4). Approxi- since 2016. The bachelor group consisted of 5 stallions of mately 50 min after the start of our observation, the oldest various ages (3 3-year-old stallions, one 4-year-old stallion, mare of the Tsetsen harem led the harem away from the and one 7-year-old stallion). The GGBSPA administration bachelors. Tsetsen followed his harem; however, he stopped confirmed that the two oldest bachelor stallions were cousins in the middle between his harem and the bachelor group (their mothers were sisters) of Tsetsen and had previously when his harem stopped. The 2-year-old stallion from the been together with Tsetsen in a bachelor group. However, Tsetsen harem did not follow the harem but remained with each of these stallions had been born into a different harem the bachelor group and rested with them while the Tsetsen (GGBSPA administration, personal communication). harem returned to Tsetsen. Afterwards, both groups (Tsetsen The observation started on 18.07.2018 at 11.29; there with his harem and the bachelor group with the 2-year-old were no water points or prime forage areas at a close dis- stallion) rested approximately 80 m apart. Approximately tance (at least 5 km from the observation point). Two other 95 min after the start of our observation, the bachelor group harems were observed from our position (∼ 1 km from the with the 2-year-old stallion from the Tsetsen harem started observation point). No interaction between them and the running towards the Tsetsen harem. At first, Tsetsen tried to Tsetsen harem or between them and the bachelor group was chase the bachelors away from his harem and the 2-year-old observed. At the beginning of the observation, all mem- stallion, but after a moment, he stopped, showed no more bers of the Tsetsen harem were resting. The bachelor group aggression, and moved back towards his harem, followed started approaching the Tsetsen harem and stopped relatively by the 2-year-old stallion from his harem and the bache- close to the Tsetsen harem (∼ 40 m). Members of the Tset- lors. Both groups merged again. When the groups split (∼ sen harem became alert, looking toward the approaching 10–15 m apart), Tsetsen remained with the bachelors, and bachelor group. After approximately 30 s, the harem stallion the 2-year-old stallion remained with the Tsetsen harem. Tsetsen moved a bit forward (∼ 5 m) in front of his harem Again, play behaviour was performed (nipping, slight rear- in the direction of the bachelor group, standing in an alert ing) between Tsetsen and three members of the bachelor position facing the bachelors. All the other members of the group (video 5). Approximately 104 min after the start of our Tsetsen harem stood close to each other in a relaxed position, observation, the 2-year-old stallion from the Tsetsen harem and none seemed to be alert anymore. After approximately moved towards the bachelor group (and Tsetsen). He was 1 min, Tsetsen started running towards the bachelors, joined followed by the Tsetsen harem, except for the oldest mare them, and the horses started to investigate each other mutu- and her 1-year-old daughter. Both groups merged again, ally. The rest of Tsetsen’s harem remained resting, paying and we observed aggressive mutual encounters between the almost no interest in the interactions between Tsetsen and mares of the Tsetsen harem and the members of the bach- the bachelors (video 1). After approximately 1 min, Tsetsen elor group, started by a kick from the 2-year-old mare from returned to his harem, and both groups stood approximately the Tsetsen harem towards the centre of the merged groups. 1 3 acta ethologica This lasted approximately 1 min, and all the members of Goldschmidt-Rothschild 2010). However, the subject of the Tsetsen harem moved back to join the oldest mare and harem holders possibly playing with members of a bach- her daughter, except for Tsetsen. He was standing with the elor group solely appears in the literature as short mentions bachelors, pawing the ground. Afterwards, he started mov- supplementing other topics (King et al. 2016; McDonnell ing towards his harem and again stopped between the two and Poulin 2002). Only Berger (1986) described the play groups (video 6). Both groups merged again after Tsetsen between harem stallions and bachelor groups of feral horses started moving back towards the bachelor groups (∼ 108 min in more detail. However, he claimed that this behaviour is after the start of our observation) and was followed by his a type of parental investment. The bachelors who played harem (video 7). After approximately 3 min, Tsetsen herded with the stallion were usually either sons of the stallion or his harem to make the harem move a bit far from the bach- grew up in his harem (they were foals when he took over the elor group (∼ 5 m). One of the adult mares kicked Tsetsen harem). The play between them happened 18 months after (video 8), the whole Tsetsen harem moved towards the bach- the bachelors left the stallions’ harem. After this time, he elor group, and both groups merged again, with no more probably did not perceive them as sons anymore and did not aggression observed (video 9). Approximately 114 min after tolerate their presence in the harem (Berger 1986). On the the start of our observation, the oldest mare of the Tset- contrary, the bachelors in our observation were not sons of sen harem started leading the whole harem away from the the harem stallion and did not grow up in his harem. Despite bachelors. The Tsetsen harem stopped approximately 20 m this fact, they were not only tolerated by the stallion, but they from the bachelor groups. We observed both groups stand- also engaged in mutual play. ing and resting. Approximately 118 min after the start of We observed a young stallion (7 years old) who became a our observation, two members of the bachelor group started harem holder for the first time in his life. Tsetsen established to walk away slowly. In contrast, the rest of the bachelor his harem by taking over half of the mares from his father’s group (3 horses) remained. Both groups were resting apart harem. One year after the described observation (in 2019), from each other (∼ 20–30 m), and no more interactions were we recorded that one 3-year-old stallion in the Tsetsen harem observed between the two groups (video 10). We remained (the then 2-year-old described interacting with the bachelor in the place until all the members of the bachelor group left group) was repeatedly mating with mares of the harem (i.e. the area and went out of sight (∼ 180 min after the start of both Tsetsen and this stallion were mating with the mares our observation). No more interactions between the Tsetsen during this breeding season). Thus, we conclude that those harem and the bachelor groups were observed by us or by two stallions formed an alliance that year (Boyd et al. 2016). the rangers after this event. The harem stallion Tsetsen was probably not the father of As a context for this observation, we observed 429 play any of the foals in the GGBSPA at the time of our observation interactions among 9 harems during the entire study period, (GGBSPA administration, personal communication). There- and just 22 of them (5%) involved the harem holder stal- fore, we conclude that the play behaviour between him and the lion. Three of the 9 harems were composed exclusively of bachelor stallions should not be attributed to parental invest- the harem holder and mares (no other stallions were in the ment, as suggested by Berger (1986). However, it is important harem). The 6 other harems that included other stallions to state that due to the population’s size and type (wild horses) further than the harem holder displayed 274 play behaviours, and the area, the reproduction records of the GGBSPA are 11 of them (4%) involving the harem holder stallion, and based on direct observations (GGBSPA administration, per- just one play interaction was observed involving the harem sonal communication). Berger (1986) also described that holder (not Tsetsen) and another stallion at the harem. familiarity, not the actual genetic relatedness, is the probable cause of harem stallion tolerance and play behaviour with bachelors. It has been proven that horses have a good memory Discussion (Hanggi and Ingersoll 2009; Lansade et al. 2020) and that familiarity, not relatedness, is the main factor in social rela- This appears to be the first evidence showing the harem tions between adult horses (Mendonça et al. 2021). Bachelor stallion (Przewalski’s horse harem holder) tolerating his groups are generally less stable, and both intra- and inter-group group to “spend time” with a group of bachelors and play- play behaviours are prevalent (Feh 2005; King et al. 2016, ing himself with the bachelor stallions. Numerous studies 2022; McDonnell and Poulin 2002; Zharkikh and Andersen describing the play behaviour of young males and members 2009). The GGBSPA administration confirmed that Tsetsen of bachelor groups have been published (e.g. Christensen was in the same bachelor group as the two oldest stallions et al. 2002; McDonnell and Haviland 1995; Zharkikh and from the bachelor group, so they know each other very well. Andersen 2009). The presence and importance of play Therefore, he might have been able to assess their fighting between the harem stallion and his foals have also been ability (Rubenstein and Hack 1992) and not perceive them as emphasised (Feh 2005; Šandlová et  al. 2020; Wells and competitors (Rubenstein 1994). He probably also knew the 1 3 acta ethologica the article's Creative Commons licence and your intended use is not other three stallions from the bachelor groups as in feral and permitted by statutory regulation or exceeds the permitted use, you will wild horse populations; all the bachelors typically know each need to obtain permission directly from the copyright holder. To view a other (Feh 2005; Rubenstein and Hack 1992; Sigurjonsdottir copy of this licence, visit http://cr eativ ecommons. or g/licen ses/ b y/4.0/ . et al. 2012). Furthermore, no sexual behaviour was observed between the members of the bachelor group and the harem mares, and the References aggressive behaviours between them were most probably caused solely by the increased proximity. For this reason, Tsetsen could Asensio N, Zandonà E, Dunn JC, Cristóbal-Azkarate J (2022) Soci- use this opportunity to maintain social bonds with members of oecological correlates of social play in adult mantled howler monkeys. Anim Behav 186:219–229. https://d oi.or g/10 .10 16/j. the bachelor group and possibly also to practice and improve anbeh av. 2022. 01. 017 his fighting skills, as frequently observed in bachelor groups Bahloul K, Pereladova OB, Soldatova N, Fisenko G, Sidorenko E, (King et al. 2016, 2022; McDonnell and Poulin 2002; Zharkikh Sempéré AJ (2001) Social organisation and dispersion of intro- and Andersen 2009). 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Appl Anim Behav Sci 78(2–4):263–290. https:// doi. or g/ 10. 1016/ S0168- Publisher's Note Springer Nature remains neutral with regard to 1591(02) 00112-0 jurisdictional claims in published maps and institutional affiliations. Mendonça RS, Pinto P, Inoue S, Ringhofer M, Godinho R, Hirata S (2021) Social determinants of affiliation and cohesion in a popula- tion of feral horses. Appl Anim Behav Sci 245:105496. https:// doi. org/ 10. 1016/j. appla nim. 2021. 105496 1 3 http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png acta ethologica Springer Journals

Novel observation of play behaviour between a harem holder and a bachelor group of Przewalski’s horses in the wild

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Abstract

Przewalski’s horses live in stable nonterritorial families (harem) of one or more harem stallions, several mares, and their offspring. The harem stallion typically behaves aggressively towards any male intruder approaching the harem. Play behav - iour is frequently observed among the group members in horses. For stallions, the most common, well-known cases of play behaviour are found between members of bachelor groups (groups of young stallions) and between the harem stallion and his offspring. The play between the harem stallion and members of the bachelor groups is, on the other hand, seemingly much rarer as a description of such events in the literature is anecdotal. In this note, we present our observation conducted in the Great Gobi B Strictly Protected Area (Mongolia) and describe the play behaviour between a Przewalski’s horse harem stallion and members of a bachelor group. The observation was done as a part of broader research during which selected Przewalski’s horses’ groups were located by binoculars in daily monitoring routines and filmed from a close distance (from 150 to 800 m). Behavioural Observation Research Interactive Software (BORIS) was used to extract data from the video recordings. The observation described in this note lasted for ∼ 180 min, during which the horses engaged in three separate bouts of play and repeated inter-group association. During the whole study (241.5 h of video recordings), this was the only observation including inter-group interactions. We observed two other events during which two harems approached each other (but never merged). Our observation is the first video recording of such an event and raises the question of how preva- lent this behaviour is in the Przewalski’s horses’ restored natural populations. We conclude that this behaviour is unusual or has not been sufficiently studied in wild or feral harems. Further technological advances may help reveal more information on this topic. Keywords Equus ferus przewalskii · Gobi · Stallion · Social behaviour · Mongolia Introduction 1985; Paquette 1994). However, the fact that play is observed in many species after sexual maturity suggests that it may be Since the earliest systematic studies of behaviour, the subject relevant for animals throughout their lifetime (Asensio et al. of why animals play has been a topic of discussion (Darwin 2022; Beckel 1991; Bond and Diamond 2003). 1871; Groos 1911). Play is mainly perceived as a tool by which Adult-only social play is typically observed among group young animals acquire skills (e.g. fighting, mating, social mates and is mainly described as a mechanism for foster- behaviour) necessary for their adult life (Martin and Caro ing social networks and encouraging collaboration among individuals. Adult play may provide a social bonding pur- pose similar to grooming, allowing animals to socialise. In other words, it enables animals to create and sustain social * Francisco Ceacero ceacero@ftz.czu.cz connections as well as restrain anger, lower tension, and enhance tolerance under challenging situations (Baldwin Faculty of Tropical AgriSciences, Czech University of Life and Baldwin 1973; Enomoto 1990; Ferguson and Frankis Sciences Prague, Prague, Czech Republic 2001; Martin and Caro 1985; Merrick 1977; Norscia and Great Gobi B Strictly Protected Area, Takhin Tal, Mongolia Palagi 2011; Palagi et al. 2006). Zoo Praha, U Trojského Zámku 120/3, Prague, Czech Republic Vol.:(0123456789) 1 3 acta ethologica In horses, play is most common and noticeable in foals stallion and members of a bachelor group in the Great Gobi and yearlings, although it also occurs in adult individu- B Strictly Protected Area (Mongolia) to raise the question of als (McDonnell and Poulin 2002). Play is also frequently how prevalent and important this behaviour is in the Przew- observed among the members of the horse bachelor groups alski’s horses’ restored natural populations and to encourage (Feh 2005), including Przewalski’s horses (Christensen et al. future research on this topic. 2002). Play behaviour is believed to be essential in devel- oping specific social skills (e.g. fighting) in young males compared to young females (Crowell-Davis et al. 1987; Rho Methods and results et al. 2007). Indeed, it has been observed that colts (male foals) interact more actively with the harem stallion (their This observation was conducted in July 2018 in the Great father) than fillies (female foals; Crowell-Davis et al. 1987; Gobi B Strictly Protected Area (GGBSPA), Mongolia, dur- Šandlová et al. 2020). However, in adult stallions with their ing broader research fieldwork (July 2018, May–June 2019, harem, observations of play with other mature stallions are and September–October 2019) focusing on the behavioural very scarce (Berger 1986; McDonnell and Poulin 2002). ecology of Przewalski’s horses. In the summer of 2018, two Przewalski’s horses (Equus ferus przewalskii) live in stable, wild-born harems involving 21 adult horses were recorded. nonterritorial family units called harems (Bourjade et al. 2009) In spring 2019, three groups were recorded (two wild-born and exhibit female defence polygyny. The harems consist of harems and one harem consisting of long-term reintroduced one or more harem stallions (males), several mares (females), mares and a wild-born stallion) involving 25 adult horses. and their young progeny (King et al. 2015; Rubenstein 2011). In autumn 2019, four groups were recorded (two wild-born When a colt or filly reaches reproductive age, typically between harems and two harems consisting of long-term reintroduced the ages of one and four, they separate from their family. While mares and wild-born stallions) involving 27 adult horses colts usually join a bachelor group until they can form their (Bernátková et al. 2022). own harem, fillies typically join another stable family unit During the whole study (241.5  h of recordings), we (Bahloul et al. 2001; Boyd and Bandi 2002; International observed only one inter-group interaction (the one described Takhi Group 2021; King et al. 2015). Mature stallions form a in this short note). Once, two harems were observed feeding new harem by taking over an existing harem or joining females close to each other (∼ 50 m) but never merged or interacted dispersed from other harems. These stallions become harem (one of these harems was Tsetsen, the one also involved in holders and protect the mares and their offspring from other the observation described here). Two harems approaching stallions and predators (Boyd et al. 2016). To guard the harem each other (neither of which was Tsetsen) were also observed successfully, the stallion typically stays more vigilant, feeds just once, but both groups stopped ∼ 300 m from each other less, moves more, and shows less behavioural synchrony than except for the harem stallions, who kept approaching each other harem members (King et al. 2016; Souris et al. 2007). other, stopped ∼ 50 m from each other, and returned to their As mares are generally protected from stallion harassment by group. the group structure, they are often in better condition, and their The GGBSPA, in SW Mongolia, is a reintroduction site reproductive success is higher than in mares not bonded to a for Przewalski’s horse and an important refuge for several breeding group (Linklater et al. 1999). It was also hypothesised other endangered species (International Takhi Group 2021; that affiliative interactions between mares and other mares or Kaczensky et al. 2008, 2011). Since 2019, it has encom- stallions might boost reproductive success by minimising har- passed ≈18 000 km of desert steppe and desert habitat assment (Cameron et al. 2009; Linklater et al. 1999). Cases (≈9 000 km before the enlargement, International Takhi of two or more stallions defending the same harem have been Group 2021). At the time of the research, there were 349 reported (Linklater et al. 2013). Previously, it was suggested Przewalski’s horses in 24 harems and 3–5 bachelor groups. that these stallion alliances improve the ability of stallions to The structure and number of the bachelor groups change defend their mutual family group from other stallions (Boyd frequently. During this research, the GGBSPA population et al. 2016) and might increase the foal survival rate in their comprised a maximum of 200 females and 149 males (GGB- group (Feh 1999). However, removal experiments showed that SPA administration, personal communication). Fieldwork the cooperative hypotheses do not appear to explain the exist- consisted of locating Przewalski’s horses’ groups with bin- ence of multi-stallion groups (Linklater et al. 2013). oculars during daily monitoring routines. Once one of the It has been described that in free-roaming populations target groups was localised, it was approached and filmed of horses, most stallions know each other and their rela- from a close distance without disturbing them (from 150 tive fighting ability (Rubenstein and Hack 1992), and most to 800 m) by a 4 K Panasonic VX1 video camera with a encounters result in ritualised behaviours (Sigurjonsdottir tripod. The ethogram consisting of 5 categories (feeding, et al. 2012). In this note, we provide a recording and descrip- locomotion, resting, social, and other) was used to catego- tion of the play behaviour between Przewalski’s horse harem rise observed behaviours. Play behaviour was included in 1 3 acta ethologica the social category (Bernátková et al. 2022). Behavioural 20 m from each other. Eighteen minutes after our observa- Observation Research Interactive Software (BORIS) was tion started, Tsetsen joined the bachelor group again. We used for event logging, coding, and observation of the cap- observed mutual investigation and play behaviour (muzzle tured videos (Friard and Gamba 2016). sniffing, slight rearing, head resting) between him and the Play between harem stallions and members of bachelor stallions of the bachelor group. No direct aggression was groups has never been observed in the population of the observed (video 2). Twenty minutes after our observation GGBSPA (GGBSPA administration, personal commu- started, one 2-year-old stallion from the Tsetsen harem nication). While observing one of the selected groups (a approached the bachelor group and Tsetsen. The rest of wild-born harem called Tsetsen harem), we noticed the non- the harem followed him. Both groups merged momentar- agonistic social behaviour of the harem stallion towards a ily, but after a few mutual aggressive encounters started bachelor group which approached his harem. The Tsetsen by the harem mares (kick threats, kicking), they split. At harem consisted of 8 individuals: 1 harem stallion (Tset- this moment, Tsetsen herded his harem and then chased the sen, 7-year-old), three adult mares (2-, 5-, and 6-year-old), bachelors away. However, 25 min after the start of our obser- 2 2-year-old males, 1 1-year-old male, and 1 1-year-old vation, the Tsetsen harem and bachelor group merged again, female. The GGBSPA administration confirmed that none of and no aggressive behaviour, herding, or any other behaviour the mares was in oestrus during the observation. Tsetsen was aiming to prevent this from the Tsetsen side was observed a harem holder for the first time in his life, and the harem this time (video 3). We observed both groups merging and was relatively new (it was formed just two months before our splitting away multiple times. During these events, Tsetsen observation). All the breeding mares remained there until mostly remained closer to the bachelor group than to his our following fieldwork one year later. Tsetsen established harem. The 2-year-old stallion from the Tsetsen harem was his harem by taking over half of the mares from his father’s also observed joining the bachelor group and Tsetsen in the harem, in which all the mares of our study had been together mutual investigation and play behaviour (video 4). Approxi- since 2016. The bachelor group consisted of 5 stallions of mately 50 min after the start of our observation, the oldest various ages (3 3-year-old stallions, one 4-year-old stallion, mare of the Tsetsen harem led the harem away from the and one 7-year-old stallion). The GGBSPA administration bachelors. Tsetsen followed his harem; however, he stopped confirmed that the two oldest bachelor stallions were cousins in the middle between his harem and the bachelor group (their mothers were sisters) of Tsetsen and had previously when his harem stopped. The 2-year-old stallion from the been together with Tsetsen in a bachelor group. However, Tsetsen harem did not follow the harem but remained with each of these stallions had been born into a different harem the bachelor group and rested with them while the Tsetsen (GGBSPA administration, personal communication). harem returned to Tsetsen. Afterwards, both groups (Tsetsen The observation started on 18.07.2018 at 11.29; there with his harem and the bachelor group with the 2-year-old were no water points or prime forage areas at a close dis- stallion) rested approximately 80 m apart. Approximately tance (at least 5 km from the observation point). Two other 95 min after the start of our observation, the bachelor group harems were observed from our position (∼ 1 km from the with the 2-year-old stallion from the Tsetsen harem started observation point). No interaction between them and the running towards the Tsetsen harem. At first, Tsetsen tried to Tsetsen harem or between them and the bachelor group was chase the bachelors away from his harem and the 2-year-old observed. At the beginning of the observation, all mem- stallion, but after a moment, he stopped, showed no more bers of the Tsetsen harem were resting. The bachelor group aggression, and moved back towards his harem, followed started approaching the Tsetsen harem and stopped relatively by the 2-year-old stallion from his harem and the bache- close to the Tsetsen harem (∼ 40 m). Members of the Tset- lors. Both groups merged again. When the groups split (∼ sen harem became alert, looking toward the approaching 10–15 m apart), Tsetsen remained with the bachelors, and bachelor group. After approximately 30 s, the harem stallion the 2-year-old stallion remained with the Tsetsen harem. Tsetsen moved a bit forward (∼ 5 m) in front of his harem Again, play behaviour was performed (nipping, slight rear- in the direction of the bachelor group, standing in an alert ing) between Tsetsen and three members of the bachelor position facing the bachelors. All the other members of the group (video 5). Approximately 104 min after the start of our Tsetsen harem stood close to each other in a relaxed position, observation, the 2-year-old stallion from the Tsetsen harem and none seemed to be alert anymore. After approximately moved towards the bachelor group (and Tsetsen). He was 1 min, Tsetsen started running towards the bachelors, joined followed by the Tsetsen harem, except for the oldest mare them, and the horses started to investigate each other mutu- and her 1-year-old daughter. Both groups merged again, ally. The rest of Tsetsen’s harem remained resting, paying and we observed aggressive mutual encounters between the almost no interest in the interactions between Tsetsen and mares of the Tsetsen harem and the members of the bach- the bachelors (video 1). After approximately 1 min, Tsetsen elor group, started by a kick from the 2-year-old mare from returned to his harem, and both groups stood approximately the Tsetsen harem towards the centre of the merged groups. 1 3 acta ethologica This lasted approximately 1 min, and all the members of Goldschmidt-Rothschild 2010). However, the subject of the Tsetsen harem moved back to join the oldest mare and harem holders possibly playing with members of a bach- her daughter, except for Tsetsen. He was standing with the elor group solely appears in the literature as short mentions bachelors, pawing the ground. Afterwards, he started mov- supplementing other topics (King et al. 2016; McDonnell ing towards his harem and again stopped between the two and Poulin 2002). Only Berger (1986) described the play groups (video 6). Both groups merged again after Tsetsen between harem stallions and bachelor groups of feral horses started moving back towards the bachelor groups (∼ 108 min in more detail. However, he claimed that this behaviour is after the start of our observation) and was followed by his a type of parental investment. The bachelors who played harem (video 7). After approximately 3 min, Tsetsen herded with the stallion were usually either sons of the stallion or his harem to make the harem move a bit far from the bach- grew up in his harem (they were foals when he took over the elor group (∼ 5 m). One of the adult mares kicked Tsetsen harem). The play between them happened 18 months after (video 8), the whole Tsetsen harem moved towards the bach- the bachelors left the stallions’ harem. After this time, he elor group, and both groups merged again, with no more probably did not perceive them as sons anymore and did not aggression observed (video 9). Approximately 114 min after tolerate their presence in the harem (Berger 1986). On the the start of our observation, the oldest mare of the Tset- contrary, the bachelors in our observation were not sons of sen harem started leading the whole harem away from the the harem stallion and did not grow up in his harem. Despite bachelors. The Tsetsen harem stopped approximately 20 m this fact, they were not only tolerated by the stallion, but they from the bachelor groups. We observed both groups stand- also engaged in mutual play. ing and resting. Approximately 118 min after the start of We observed a young stallion (7 years old) who became a our observation, two members of the bachelor group started harem holder for the first time in his life. Tsetsen established to walk away slowly. In contrast, the rest of the bachelor his harem by taking over half of the mares from his father’s group (3 horses) remained. Both groups were resting apart harem. One year after the described observation (in 2019), from each other (∼ 20–30 m), and no more interactions were we recorded that one 3-year-old stallion in the Tsetsen harem observed between the two groups (video 10). We remained (the then 2-year-old described interacting with the bachelor in the place until all the members of the bachelor group left group) was repeatedly mating with mares of the harem (i.e. the area and went out of sight (∼ 180 min after the start of both Tsetsen and this stallion were mating with the mares our observation). No more interactions between the Tsetsen during this breeding season). Thus, we conclude that those harem and the bachelor groups were observed by us or by two stallions formed an alliance that year (Boyd et al. 2016). the rangers after this event. The harem stallion Tsetsen was probably not the father of As a context for this observation, we observed 429 play any of the foals in the GGBSPA at the time of our observation interactions among 9 harems during the entire study period, (GGBSPA administration, personal communication). There- and just 22 of them (5%) involved the harem holder stal- fore, we conclude that the play behaviour between him and the lion. Three of the 9 harems were composed exclusively of bachelor stallions should not be attributed to parental invest- the harem holder and mares (no other stallions were in the ment, as suggested by Berger (1986). However, it is important harem). The 6 other harems that included other stallions to state that due to the population’s size and type (wild horses) further than the harem holder displayed 274 play behaviours, and the area, the reproduction records of the GGBSPA are 11 of them (4%) involving the harem holder stallion, and based on direct observations (GGBSPA administration, per- just one play interaction was observed involving the harem sonal communication). Berger (1986) also described that holder (not Tsetsen) and another stallion at the harem. familiarity, not the actual genetic relatedness, is the probable cause of harem stallion tolerance and play behaviour with bachelors. It has been proven that horses have a good memory Discussion (Hanggi and Ingersoll 2009; Lansade et al. 2020) and that familiarity, not relatedness, is the main factor in social rela- This appears to be the first evidence showing the harem tions between adult horses (Mendonça et al. 2021). Bachelor stallion (Przewalski’s horse harem holder) tolerating his groups are generally less stable, and both intra- and inter-group group to “spend time” with a group of bachelors and play- play behaviours are prevalent (Feh 2005; King et al. 2016, ing himself with the bachelor stallions. Numerous studies 2022; McDonnell and Poulin 2002; Zharkikh and Andersen describing the play behaviour of young males and members 2009). The GGBSPA administration confirmed that Tsetsen of bachelor groups have been published (e.g. Christensen was in the same bachelor group as the two oldest stallions et al. 2002; McDonnell and Haviland 1995; Zharkikh and from the bachelor group, so they know each other very well. Andersen 2009). The presence and importance of play Therefore, he might have been able to assess their fighting between the harem stallion and his foals have also been ability (Rubenstein and Hack 1992) and not perceive them as emphasised (Feh 2005; Šandlová et  al. 2020; Wells and competitors (Rubenstein 1994). He probably also knew the 1 3 acta ethologica the article's Creative Commons licence and your intended use is not other three stallions from the bachelor groups as in feral and permitted by statutory regulation or exceeds the permitted use, you will wild horse populations; all the bachelors typically know each need to obtain permission directly from the copyright holder. To view a other (Feh 2005; Rubenstein and Hack 1992; Sigurjonsdottir copy of this licence, visit http://cr eativ ecommons. or g/licen ses/ b y/4.0/ . et al. 2012). Furthermore, no sexual behaviour was observed between the members of the bachelor group and the harem mares, and the References aggressive behaviours between them were most probably caused solely by the increased proximity. For this reason, Tsetsen could Asensio N, Zandonà E, Dunn JC, Cristóbal-Azkarate J (2022) Soci- use this opportunity to maintain social bonds with members of oecological correlates of social play in adult mantled howler monkeys. Anim Behav 186:219–229. https://d oi.or g/10 .10 16/j. the bachelor group and possibly also to practice and improve anbeh av. 2022. 01. 017 his fighting skills, as frequently observed in bachelor groups Bahloul K, Pereladova OB, Soldatova N, Fisenko G, Sidorenko E, (King et al. 2016, 2022; McDonnell and Poulin 2002; Zharkikh Sempéré AJ (2001) Social organisation and dispersion of intro- and Andersen 2009). Another reason for Tsetsen’s tolerance duced kulans (Equus hemionus kulan) and Przewalski horses (Equus przewalski) in the Bukhara reserve. Uzbekistan Journal and “friendliness” might be that he was an inexperienced harem of Arid Environments 47(3):309–323. https:// doi. org/ 10. 1006/ holder (Boyd et al. 2016) and had limited control over his harem JARE. 2000. 0714 (Linklater et al. 2013), as he seemingly formed an alliance with a Baldwin JD, Baldwin JI (1973) The role of play in social organisa- 3-year-old stallion (his half-brother) from his harem 1 year after tion: comparative observations on squirrel monkeys (Saimiri). Primates 14(4):369–381. https:// doi. org/ 10. 1007/ BF017 31358 this observation was conducted (Boyd et al. 2016). Beckel AL (1991) Wrestling play in adult river otters, Lutra canaden- In summary, our observation appears to be the first video sis. 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Journal

acta ethologica – Springer Journals

Published: Jun 1, 2023

Keywords: Equus ferus przewalskii; Gobi; Stallion; Social behaviour; Mongolia

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