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Leadership is a risky behaviour that can impact individuals and groups. Leaders, i.e. individuals who perform or initiate a task while other individuals in the group follow, have been studied in different contexts, but there is still a lack of understanding on the role of individual characteristics that may predispose them to become leaders, such as dominance and personality. In particular, the characteristics of leaders in domestic animal populations has been poorly examined. We studied leadership within 32 groups of young pigs (Sus scrofa domesticus, n = 366 individuals). Leadership was assessed during a group-based fear test (Human Approach Test) which was repeated three times. The first individual per group to touch the person was identified as leader. We assessed repeatability of leadership and characteristics of leaders as compared to followers. Leadership was marginally repeatable, with 6 out of 26 groups having a consistent single leader across all tests. Females had odds 4.13 times greater than males of being a leader, while there was no effect of body weight (a proxy of dominance) or coping style on leadership. The results indicate a similarity with wild populations, in which females lead the herd even though the males, which are superior in body weight, are often dominant.

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In recent decades, the integration of horses (Equus ferus) in European rewilding initiatives has gained widespread popularity due to their potential for regulating vegetation and restoring natural ecosystems. However, employing horses in conservation efforts presents important challenges, which we here explore and discuss. These challenges encompass the lack of consensus on key terms inherent to conservation and rewilding, the entrenched culture and strong emotions associated with horses, low genetic diversity and high susceptibility to hereditary diseases in animals under human selection, as well as insufficient consideration for the social behaviour of horses in wild-living populations. In addition, management of wild-living horses involves intricate welfare, ethics and legislative dimensions. Anthropocentric population-control initiatives may be detrimental to horse group structures since they tend to prioritise individual welfare over the health of populations and ecosystems. To overcome these challenges, we provide comprehensive recommendations. These involve a systematic acquisition of genetic information, a focus on genetic diversity rather than breed purity and minimal veterinary intervention in wild-living populations. Further, we advise allowing for natural top-down and bottom-up control - or, if impossible, simulating this by culling or non-lethal removal of horses - instead of using fertility control for population management. We advocate for intensified collaboration between conservation biologists and practitioners and enhanced communication with the general public. Decision-making should be informed by a thorough understanding of the genetic makeup, common health issues and dynamics, and social behaviour in wild-living horse populations. Such a holistic approach is essential to reconcile human emotions associated with horses with the implementation of conservation practices that are not only effective but also sustainable for the long-term viability of functional, biodiverse ecosystems, while rehabilitating the horse as a widespread wild-living species in Europe.

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All animals exhibit some combination of spatial and social behaviours. A diversity of interactions occurs between such behaviours, producing emergent phenomena at the spatial-social interface. Untangling and interrogating these complex, intertwined processes can be vital for identifying the mechanisms, causes and consequences of behavioural variation in animal ecology. Nevertheless, the integrated study of the interactions between spatial and social phenotypes and environments (at the spatial-social interface) is in its relative infancy. In this theme issue, we present a collection of papers chosen to expand the spatial-social interface along several theoretical, methodological and empirical dimensions. They detail new perspectives, methods, study systems and more, as well as offering roadmaps for applied outputs and detailing exciting new directions for the field to move in the future. In this Introduction, we outline the contents of these papers, placing them in the context of what comes before, and we synthesize a number of takeaways and future directions for the spatial-social interface. This article is part of the theme issue 'The spatial-social interface: a theoretical and empirical integration'.

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Animals simultaneously navigate spatial and social environments, and their decision-making with respect to those environments constitutes their spatial (e.g. habitat selection) and social (e.g. conspecific associations) phenotypes. The spatial-social interface is a recently introduced conceptual framework linking these components of spatial and social ecology. The spatial-social interface is inherently scale-dependent, yet it has not been integrated with the rich body of literature on ecological scale. Here, we develop a conceptual connection between the spatial-social interface and ecological scale. We propose three key innovations that incrementally build upon each other. First, the use-availability framework that underpins a large body of literature in behavioural ecology can be used in analogy to the phenotype-environment nomenclature and is transferable across the spatial and social realms. Second, both spatial and social phenotypes are hierarchical, with nested components that are linked via constraints-from the top down-or emergent properties-from the bottom up. Finally, in both the spatial and social realms, the definitions of environment and phenotype depend on the focal scale of inquiry. These conceptual innovations cast our understanding of the relationships between social and spatial dimensions of animal ecology in a new light, allowing a more holistic understanding and clearer hypothesis development for animal behaviour. This article is part of the theme issue 'The spatial-social interface: a theoretical and empirical integration'.

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Social isolation causes profound changes in social behaviour in a variety of species. However, the genetic and molecular mechanisms modulating behavioural responses to social isolation and social recovery remain to be elucidated. Here, we quantified the behavioural response of vinegar flies to social isolation using two distinct protocols (social space preference and sociability, the spontaneous tendencies to form groups). We found that social isolation increased social space and reduced sociability. These effects of social isolation were reversible and could be reduced after 3 days of group housing. Flies with a loss of function of neuroligin3 (orthologue of autism-related neuroligin genes) with known increased social space in a socially enriched environment were still able to recover from social isolation. We also show that dopamine (DA) is needed for a response to social isolation and recovery in males but not in females. Furthermore, only in males, DA levels are reduced after isolation and are not recovered after group housing. Finally, in socially enriched flies mutant for neuroligin3, DA levels are reduced in males, but not in females. We propose a model to explain how DA and neuroligin3 are involved in the behavioural response to social isolation and its recovery in a dynamic and sex-specific manner.

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Fluctuations in estradiol levels at each stage of life in women are considered one of the causes of mental diseases through their effects on the central nervous system. During menopause, a decrease in estradiol levels has been reported to affect the serotonin nervous system and induce depression-like and anxiety symptoms. However, the regulation of brain and behaviour during childhood and adolescence is poorly understood. Moreover, the role of oestrogen receptors α and ß in the regulation of the serotonergic nervous system has been reported, but little is known about the involvement of G protein-coupled receptor 30. Therefore, in this study, we used an ovariectomized childhood mouse model to analyse behaviour and investigate the effects on the serotonin nervous system. We showed that ovariectomy surgery at 4 weeks of age, which is the weaning period, induced a decrease in spontaneous locomotor activity during the active period and a preference for novel mice over familiar mice in the three-chamber social test at 10 weeks of age. In addition, the administration of G-1, a protein-coupled receptor 30 agonist, to ovariectomized mice suppressed spontaneous locomotor activity and the preference for novel mice. Furthermore, we demonstrated that childhood ovariectomy induces increased tryptophan hydroxylase gene expression in the raphe nucleus and increased serotonin release in the amygdaloid nucleus, and administration of G-1 ameliorated these effects. Our study suggests that G protein-coupled receptor 30-mediated regulation of serotonin synthesis is involved in changes in activity and social-cognitive behaviour due to decreased estradiol levels during childhood.

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Young precocial birds benefit from staying close to both their mother and their siblings, while prioritising adults, which provide better care. Which features of the stimuli are used by young birds to prioritise approach and eventually attachment to adults over siblings is unknown. We started to address this question in newly hatched domestic chicks (Gallus gallus), focusing on their spontaneous preferences for visual features that systematically vary between adult and juvenile chickens, and that had previously been identified as attractive: size (larger in adults than in juveniles) and colour (darker and redder in adults than in juveniles). Overall, chicks at their first visual experience, that had never seen a conspecific beforehand, were most attracted to the red and large stimuli (two adult features) and spent more time in close proximity with red stimuli than with yellow stimuli. When tested with red large versus small objects (Exp. 1), chicks preferred the large shape. When tested with yellow large and small objects (Exp. 2), chicks did not show a preference. Chicks had a stronger preference for large red stimuli (vs. small yellow objects) than for small red stimuli (vs. a large yellow object) (Exp. 3). These results suggest that the combination of size and colour form the predisposition that helps chicks to spontaneously discriminate between adult and juvenile features from the first stages of life, in the absence of previous experience, exhibiting a preference to approach stimuli with features associated with the presence of adult conspecifics.

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Autism spectrum disorder (ASD) has proven to be highly enigmatic due to the diversity of its underlying genetic causes and the huge variability in symptom presentation. Uncovering common phenotypes across people with ASD and pre-clinical models allows us to better understand the influence on brain function of the many different genetic and cellular processes thought to contribute to ASD aetiology. One such feature of ASD is the convergent evidence implicating abnormal functioning of the medial prefrontal cortex (mPFC) across studies. The mPFC is a key part of the 'social brain' and may contribute to many of the changes in social behaviour observed in people with ASD. Here we review recent evidence for mPFC involvement in both ASD and social behaviours. We also highlight how pre-clinical mouse models can be used to uncover important cellular and circuit-level mechanisms that may underly atypical social behaviours in ASD. This article is part of the Special Issue on "PFC circuit function in psychiatric disease and relevant models".

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This study extended the theory of planned behaviour (TPB) and applied it to analyse influencing factors of food waste separation intention (FSI) among households in central Vietnam. Partial least squares structural equation modelling (PLS-SEM) was employed to analyse the data. The heterogeneity of factors contributing to FSI in cities of three levels was examined using multi-group analysis (MGA). The results indicate that attitudes, subjective norms (SN), perceived behavioural control (PBC), awareness of benefit (AB), information publicity (IP), facility availability (FA), and trust significantly affected FSI of households. MGA results indicated the heterogeneity of impacts of PBC and attitude on FSI of households among three municipal levels. The results will serve as basic data for waste officers, solid-waste management project leaders, non-governmental organisations (NGOs), and other related stakeholders to lay the foundation of food waste management planning in terms of regional scale and local scale. This study will also aid the creation of a circular economy by providing a scientific base for enhancing food waste separation at source in central Vietnam.

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BACKGROUND: Social robots are promising tools to improve the quality of life of children and youth living with anxiety and should be developed based on the priorities of end users. However, pathways to include young people in patient-oriented research, particularly in the overlap between technology and mental health, have been historically limited. OBJECTIVE: In this work, we describe engagement with experts with lived experiences of paediatric anxiety in a social robotics research programme. We report the experiences of patient advisors in a co-creation process and identify considerations for other research groups looking to involve end users in technology development in the field of youth mental health. DESIGN: We engaged individuals with a lived experience of paediatric anxiety (current, recent past, or from a parent perspective) using three different models over the course of three years. Two initial patient partners were involved during project development, eight were engaged as part of an advisory panel ('the League') during study development and data analysis and four contributed as ongoing collaborators in an advisory role. League members completed a preparticipation expectation survey and a postparticipation experience survey. FINDINGS: Eight individuals from a range of anxiety-related diagnostic groups participated in the League as patient partners. Members were teenagers (n = 3), young adults aged 22-26 years who had connected with a youth mental health service as children within the past eight years (n = 3) or parents of children presently living with anxiety (n = 2). Preferred methods of communication, expectations and reasons for participating were collected. The League provided specific and actionable feedback on the design of workshops on the topic of social robotics, which was implemented. They reported that their experiences were positive and fairly compensated, but communication and sustained engagement over time were challenges. Issues of ethics and language related to patient-centred brain health technology research are discussed. CONCLUSIONS: There is an ethical imperative to meaningfully incorporate the voices of youth and young adults with psychiatric conditions in the development of devices intended to support their mental health and quality of life. PATIENT OR PUBLIC CONTRIBUTION: Six young people and two parents with lived experiences of paediatric anxiety participated in all stages of developing a research programme on social robotics to support paediatric mental health in a community context. They also provided input during the preparation of this manuscript.

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The brain's sensory lateralization involves the processing of information from the sensory organs primarily in one hemisphere. This can improve brain efficiency by reducing interference and duplication of neural circuits. For species that rely on successful interaction among family partners, such as geese, lateralization can be advantageous. However, at the group level, one-sided biases in sensory lateralization can make individuals predictable to competitors and predators. We investigated lateral preferences in the positioning of pair mates of Greater white-fronted geese Anser albifrons albifrons. Using GPS-GSM trackers, we monitored individual geese in flight throughout the year. Our findings indicate that geese exhibit individual lateral biases when viewing their mate in flight, but the direction of these biases varies among individuals. We suggest that these patterns of visual lateralization could be an adaptive trait for the species with long-term social monogamy, high levels of interspecies communication and competition, and high levels of predator and hunting pressure.

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BACKGROUND: Social behaviour plays a key role in mental health and wellbeing, and developing greater understanding of mechanisms underlying social interaction-particularly social motivation-holds substantial transdiagnostic impact. Common rodent behavioural assays used to assess social behaviour are limited in their assessment of social motivation, whereas the social operant conditioning model can provide unique and valuable insights into social motivation. Further characterisation of common experimental parameters that may influence social motivation within the social operant model, as well as complementary methodological and analytical approaches, are warranted. METHODS: This study investigated the effects of biological sex, housing condition, and time-of-day, on social motivation using the social operant model. This involved training rats to lever press (FR1) for 60-s access to a social reward (same-sex conspecific stimulus). Subjects were male and female Wistar rats, housed under individual or paired conditions, and sessions were conducted either in the mid-late light phase (ZT6-10) or early-mid dark phase (ZT13-17). A behavioural economics approach was implemented to measure social demand and the influence of stimulus partner sex (same- vs. opposite-sex stimulus) on social operant responding. Additionally, video tracking analyses were conducted to assess the degree of convergence between social appetitive and consummatory behaviours. RESULTS: Biological sex, housing conditions, the interaction between sex and housing, and stimulus partner sex potently influenced social motivation, whereas time-of-day did not. Behavioural economics demonstrated that sex, housing, and their interaction influence both the hedonic set-point and elasticity of social demand. Video analysis of social interaction during social operant sessions revealed that social appetitive and consummatory behaviours are not necessarily convergent, and indicate potential social satiety. Lastly, oestrus phase of female experimental and stimulus rats did not impact social motivation within the model. CONCLUSIONS: Social isolation-dependent sex differences exist in social motivation for rats, as assessed by social operant conditioning. The social operant model represents an optimal preclinical assay that comprehensively evaluates social motivation and offers a platform for future investigations of neurobiological mechanisms underlying sex differences in social motivation. These findings highlight the importance of continued consideration and inclusion of sex as a biological variable in future social operant conditioning studies. Humans are social creatures-our everyday interactions with others and the support this provides play a key role in our wellbeing. For those experiencing mental health conditions, people's motivation to engage with others can wane, which can lead them to withdraw from those who support them. Therefore, to develop better treatment strategies for these conditions, we need to gain a deeper understanding of social motivation. Studying social behaviour in animals can facilitate this investigation of social motivation as it allows for a causal understanding of underlying neurobiology that is not possible in human experiments. An optimal way to study social motivation in animals is using the social operant conditioning model, where rats learn to press a lever that opens a door and allows them to interact with another rat for a short time. This study characterised the social operant model by testing whether sex, housing conditions, time-of-day, and the sex of the stimulus partner influence rats' motivation to seek interaction with another rat. We found that female rats were more socially motivated than males, and that rats living alone were more motivated than those living with another rat; interestingly, this effect of housing affected females more than males. Regardless of sex, rats were more motivated to interact with a rat of the opposite sex. These findings provide insights into sex differences in social motivation in rats and new insights into the social operant model which will help guide future research into social motivation and other mental health conditions.

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There is increasing evidence that competent handling of social interactions among conspecifics has positive effects on individual fitness. While individual variation in social competence has been appreciated, the role of long-term experience in the acquisition of superior social skills has received less attention. With the goal of promoting further research, we integrate knowledge across disciplines to assess social expertise, defined as the characteristics, skills and knowledge allowing individuals with extensive social experience to perform significantly better than novices on a given social task. We focus on three categories of social behaviour. First, animals can gain from adjusting social behaviour towards individually recognised conspecifics that they interact with on a regular basis. For example, there is evidence that some territorial animals individually recognise their neighbours and modify their social interactions based on experience with each neighbour. Similarly, individuals in group-living species learn to associate with specific group members based on their expected benefits from such social connections. Individuals have also been found to devote considerable time and effort to learning about the spatial location and timing of sexual receptivity of opposite-sex neighbours to optimise reproduction. Second, signallers can enhance their signals, and receivers can refine their response to signals with experience. In many birds and insects, individuals can produce more consistent signals with experience, and females across a wide taxonomic range can adaptively adjust mating preferences after perceiving distinct male signals. Third, in many species, individuals that succeed in reproducing encounter the novel, complex task of caring for vulnerable offspring. Evidence from a few species of mammals indicates that mothers improve in providing for and protecting their young over successive broods. Finally, for social expertise to evolve, heritable variation in social expertise has to be positively associated with fitness. Heritable variation has been shown in traits contributing to social expertise including social attention, empathy, individual recognition and maternal care. There are currently limited data associating social expertise with fitness, most likely owing to sparse research effort. Exceptions include maternal care, signal refinement, and familiarity with neighbours and group members. Overall, there is evidence that individuals in many species keep refining their social skills with experience throughout life. Hence we propose promising lines of research that can quantify more thoroughly the development of social expertise and its effects on fitness.

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BACKGROUND: Oral and social problems can exacerbate long-term care. Understanding the relationship between social aspects and oral hypofunction can help identify high-risk factors for long-term care. OBJECTIVE: This study aimed to investigate the social aspects of oral hypofunction among medical outpatients. METHODS: This retrospective cross-sectional study included patients who visited an outpatient clinic for frailty. The oral function was assessed using seven items: oral hygiene, occlusal force, masticatory function, tongue-lip motor function, tongue pressure, oral dryness and swallowing function. Participants with three or more functional declines were classified as having 'oral hypofunction'. Social aspects were assessed using 21 items in four categories: general resources, basic social needs, social resources and social behaviours/activities. We analysed the relationship between oral hypofunction and each social aspect. RESULTS: A total of 316 participants (age 78.5 ± 6.4 years) were included, and 128 (41%) had oral hypofunction. Participants with oral hypofunction were significantly more likely to have limited education, require long-term care, not use transportation, depend on others for shopping, not participate in events, lack association membership and not engage in charity or volunteer work. After logistic regression analysis, 'long-term care' and 'no association membership' remained significantly associated with oral hypofunction (odds ratios 2.3, 2.3; 95% CI 1.1-5.0, 1.1-4.7, respectively). CONCLUSION: Participants with oral hypofunction faced challenges in 'general resources' and 'social behaviour/activities', which were linked to oral function issues. Future longitudinal studies are necessary to further investigate this relationship.

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Anthropogenic (man-made) noise constitutes a novel and widespread pollutant which is increasing in prevalence in terrestrial and aquatic ecosystems, resulting in alterations of natural soundscapes. There is proliferating evidence that noise leads to maladaptive behaviour in wildlife, yet few studies have addressed the effect on mammalian parent-offspring interactions. We investigated the impact of road noise on dwarf mongoose (Helogale parvula) offspring nearest-neighbour decision-making while foraging, using a field-based playback experiment. We predicted that offspring would forage closer to groupmates, especially adult and dominant individuals, when experiencing road noise compared with ambient sound to reduce communication masking and alleviate stress. We also predicted that noise would have a reduced effect with increasing offspring age owing to reduced reliance on adult groupmates for provisioning and predator defence. However, we found that mean nearest-neighbour distance and nearest-neighbour intrinsic characteristics (age, sex and dominance status) did not differ significantly between sound treatments, and these responses did not vary significantly with focal individual age. Noise may not impact nearest-neighbour decision-making owing to habituation from chronic natural exposure; alternatively, noise could induce stress and distraction, resulting in maladaptive decision-making. Future work should aim to detangle the underlying mechanisms mediating parent-offspring interactions in conditions of anthropogenic noise.

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(1) Background: The effects of short-term social isolation during adulthood have not yet been fully established in rats behaviourally, and not at all transcriptomically in the medial prefrontal cortex (mPFC). (2) Methods: We measured the behavioural effects of housing adult male rats in pairs or alone for 10 days. We also used RNA sequencing to measure the accompanying gene expression alterations in the mPFC of male rats. (3) Results: The isolated animals exhibited reduced sociability and social novelty preference, but increased social interaction. There was no change in their aggression, anxiety, or depression-like activity. Transcriptomic analysis revealed a differential expression of 46 genes between the groups. The KEGG pathway analysis showed that differentially expressed genes are involved in neuroactive ligand-receptor interactions, particularly in the dopaminergic and peptidergic systems, and addiction. Subsequent validation confirmed the decreased level of three altered genes: regulator of G protein signalling 9 (Rgs9), serotonin receptor 2c (Htr2c), and Prodynorphin (Pdyn), which are involved in dopaminergic, serotonergic, and peptidergic function, respectively. Antagonizing Htr2c confirmed its role in social novelty discrimination. (4) Conclusions: Social homeostatic regulations include monoaminergic and peptidergic systems of the mPFC.

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The word zeitgeist refers to common perceptions shared in a given culture. Meanwhile, a defining feature of loneliness is feeling that one's views are not shared with others. Does loneliness correspond with deviating from the zeitgeist? Across two independent brain imaging datasets, lonely participants' neural representations of well-known celebrities strayed from group-consensus neural representations in the medial prefrontal cortex-a region that encodes and retrieves social knowledge (Studies 1 A/1B: N = 40 each). Because communication fosters social connection by creating shared reality, we next asked whether lonelier participants' communication about well-known celebrities also deviates from the zeitgeist. Indeed, when a strong group consensus exists, lonelier individuals use idiosyncratic language to describe well-known celebrities (Study 2: N = 923). Collectively, results support lonely individuals' feeling that their views are not shared. This suggests loneliness may not only reflect impoverished relationships with specific individuals, but also feelings of disconnection from prevalently shared views of contemporary culture.

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While humans are highly cooperative, they can also behave spitefully. Yet spite remains understudied. Spite can be normatively driven and while previous experiments have found some evidence that cooperation and punishment may spread via social learning, no experiments have considered the social transmission of spiteful behaviour. Here we present an online experiment where, following an opportunity to earn wealth, we asked participants to choose an action towards an anonymous partner across a full spectrum of social behaviour, from spite to altruism. In accordance with cultural evolutionary theory, participants were presented with social information that varied in source and content. Across six conditions, we informed participants that either the majority or the highest earner had chosen to behave spitefully, neutrally or altruistically. We found an overall tendency towards altruism, but at lower levels among those exposed to spite compared with altruism. We found no difference between social information that came from the majority or the highest earner. Exploratory analysis revealed that participants' earnings negatively correlated with altruistic behaviour. Our results contrast with previous literature that report high rates of spite in experimental samples and a greater propensity for individuals to copy successful individuals over the majority.

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Cooperation is common in animals, yet the specific mechanisms driving collaborative behaviour in different species remain unclear. We investigated the proximate mechanisms underlying the cooperative behaviour of bumblebees in two different tasks, where bees had to simultaneously push a block in an arena or a door at the end of a tunnel for access to reward. In both tasks, when their partner's entry into the arena/tunnel was delayed, bees took longer to first push the block/door compared with control bees that learned to push alone. In the tunnel task, just before gaining access to reward, bees were more likely to face towards their partner than expected by chance or compared with controls. These results show that bumblebees' cooperative behaviour is not simply a by-product of individual efforts but is socially influenced. We discuss how bees' turning behaviours, e.g. turning around before first reaching the door when their partner was delayed and turning back towards the door in response to seeing their partner heading towards the door, suggest the potential for active coordination. However, because these behaviours could also be interpreted as combined responses to social and secondary reinforcement cues, future studies are needed to help clarify whether bumblebees truly use active coordination.

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Sociopositive interactions with conspecifics are essential for equine welfare and quality of life. This study aimed to validate the use of wearable ultra-wideband (UWB) technology to quantify the spatial relationships and dynamics of social behaviour in horses by continuous (1/s) measurement of interindividual distances. After testing the UWB devices' spatiotemporal accuracy in a static environment, the UWB measurement validity, feasibility and utility under dynamic field conditions was assessed in a group of 8 horses. Comparison of the proximity measurements with video surveillance data established the measurement accuracy and validity (r = 0.83, p < 0.0001) of the UWB technology. The utility for social behaviour research was demonstrated by the excellent accordance of affiliative relationships (preferred partners) identified using UWB with video observations. The horses remained a median of 5.82 m (95% CI 5.13-6.41 m) apart from each other and spent 20% (median, 95% CI 14-26%) of their time in a distance ≤ 3 m to their preferred partner. The proximity measurements and corresponding speed calculation allowed the identification of affiliative versus agonistic approaches based on differences in the approach speed and the distance and duration of the resulting proximity. Affiliative approaches were statistically significantly slower (median: 1.57 km/h, 95% CI 1.26-1.92 km/h, p = 0.0394) and resulted in greater proximity (median: 36.75 cm, 95% CI 19.5-62 cm, p = 0.0003) to the approached horse than agonistic approaches (median: 3.04 km/h, 95% CI 2.16-3.74 km/h, median proximity: 243 cm, 95% CI 130-319 cm), which caused an immediate retreat of the approached horse at a significantly greater speed (median: 3.77 km/h, 95% CI 3.52-5.85 km/h, p < 0.0001) than the approach.

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