"We forgot about the donkeys!" An institutional analysis of the shift in animal welfare from direct implementation towards advocacy-based programming.

Shifts from direct implementation to advocacy-based programming have been documented across many non-governmental organisation (NGO) sectors, including animal welfare. Semi-structured interviews with 32 staff from different positions within animal welfare NGOs explored recent programming changes. Maintaining a balance between direct implementation and advocacy-based activities emerged as a strong theme. The findings suggest that risks are associated with both the direct implementation status quo and transitioning to an advocacy-based focus. Risks of the former include treating symptoms rather than root causes of welfare problems. Organisational change can be disruptive and necessitates realignment of core competences, in turn influencing NGO mission. Identified risks of transition include loss of individuals whose values fail to align with new programming directions, increased upwards accountability requirements for accessing institutional donors and difficulties when phasing out direct implementation approaches. Whilst having to be dynamic, NGOs need to evaluate the risks associated with programming decisions, considering their vision, mission and staff identity in order to ensure that welfare programming is as effective as possible.

Belief in Animal Sentience and Affective Owner Attitudes are linked to Positive Working Equid Welfare across Six Countries.

Belief in animal sentience and the quality of human-animal relationships play a significant role in animal welfare. However, the link between an individual animal's welfare and the beliefs and emotional connection of the owner to the animal is understudied and focussed on single cultures, limiting generalisability. In this study, we explored potential links between owner attitude, beliefs in animal sentience, and working equid welfare across four continents. This study used a welfare assessment protocol alongside a questionnaire exploring owner attitudes to assess 378 participants across six countries. Overall, the general health status and body condition of equids belonging to owners with an affective (versus instrumental) perspective and those owners who believed that their equids could feel emotions, were significantly better. Equids belonging to owners who believed that their animals could feel pain were also significantly less likely to be lame. Potential causal relationships between factors and theories explaining these beliefs are discussed. These results can inform future welfare initiatives by highlighting the importance of the human-equid relationship and the role that beliefs regarding animal sentience play in influencing equid welfare.

Risk factors for stereotypic behaviour in captive ungulates.

Behavioural needs are highly motivated actions critical to a species survival and reproduction. Prolonged restriction of these behaviours can lead to stereotypic behaviours (SB) in captive animals, and this is particularly common in ungulate species. While risk factors for SB have been suggested for some ungulates, no study has integrated these findings to identify which aspects of ungulates' wild behavioural biology and captive husbandry are potential drivers for SB across this clade. We collated SB data from 15 236 individuals across 38 ungulate species from 95 sources, and determined species wild/free-ranging behaviour from 559 additional studies. Bayesian-phylogenetic statistical methods showed that ungulate behavioural needs relating to foraging and mating are particularly affected by captive environments, with promiscuous and browsing species showing the greatest prevalence of SB. Concentrate-only diets and lack of ad libitum feed substrates were also associated with high SB prevalence. This study identifies which ungulates are better suited to captive environments and which species require targeted husbandry, enrichment and breeding protocols in order to meet their behavioural needs. Our approach of applying Bayesian-phylogenetic inference to factors influencing SB within a clade can be used to identify other intrinsic and extrinsic risk factors of reduced animal health and welfare.

"What can we do to actually reach all these animals?" Evaluating approaches to improving working equid welfare.

NGOs utilise a range of programming to bring about desired changes. This study examines the advantages and limitations of the range of approaches used by one particular NGO sector: working equid welfare organisations, from the perspective of NGO staff. NGO staff hold a wealth of knowledge regarding the design and implementation of welfare improvement initiatives; however this knowledge is rarely systematically documented. Through 32 semi-structured interviews the experiences of staff across multiple organisations were explored. The nine most commonly utilised approaches within equid focussed NGO programming were discussed with participants. Many themes resulting from the interviews centred around the strengths and disadvantages of these individual approaches, implemented both historically and presently by different NGOs. The influences of the context in which initiatives took place and barriers to welfare improvement that existed across approaches were also examined. Striking a balance came through as a strong theme at many levels. Balancing top-down versus bottom-up approaches was a frequently voiced concern, as was balancing the utility of certain approaches against associated factors warranting caution. Combinations of approaches that were felt to be complementary provided a balance that drew on the strengths and mitigated for the weaknesses of different approaches. The need to tailor approaches to individual contexts was also raised and is especially relevant to INGOs working across multiple countries, cultures and political structures. The study provides an informed insight into potential factors to consider when designing future welfare initiatives. The collaborative pooling of experience across different NGOs could help make welfare initiatives more effective and provide a framework for NGOs in other fields to learn from each other's collective knowledge.

Non-random associations in group housed rats (Rattus norvegicus).

Ecological factors, such as predation, have traditionally been used to explain sociability. However, it is increasingly recognised that individuals within a group do not associate randomly, and that these non-random associations can generate fitness advantages. The majority of the empirical evidence on differentiated associations in group-living mammals, however, comes from a limited number of taxa and we still know very little about their occurrence and characteristics in some highly social species, such as rats (Rattus spp.). Here, using network analysis, we quantified association patterns in four groups of male fancy rats. We found that the associations between rats were not randomly distributed and that most individuals had significantly more preferred/avoided associates than expected by random. We also found that these preferences can be stable over time, and that they were not influenced by individuals' rank position in the dominance hierarchy. Our findings are consistent with work in other mammals, but contrast with the limited evidence available for other rat strains. While further studies in groups with different demographic composition are warranted to confirm our findings, the occurrence of differentiated associations in all male groups of rats have important implications for the management and welfare of captive rat populations.

Evaluation of long-term welfare initiatives on working equid welfare and social transmission of knowledge in Mexico.

Working equids play an essential role in supporting livelihoods, providing resilience and income security to people around the world, yet their welfare is often poor. Consequently, animal welfare focussed NGOs employ a range of initiatives aimed at improving standards of working equid welfare. However, there is debate surrounding the efficacy of welfare initiatives utilised and long term monitoring and evaluation of initiatives is rarely undertaken. This study compares equid welfare and the social transmission of welfare information across Mexican communities that had previously received differing intervention histories (veterinary treatment plus educational initiatives, veterinary treatment only and control communities) in order to assess their efficacy. Indicators of equid welfare were assessed using the Equid Assessment Research and Scoping tool and included body condition score, skin alterations, lameness, general health status and reaction to observer approach. Owners were interviewed about their involvement in previous welfare initiatives, beliefs regarding equid emotions and pain, and the social transmission of welfare knowledge, including whether they ask advice about their equid or discuss its health with others and whether there is a specific individual that they consider to be 'good with equids' in their community. In total 266 owners were interviewed from 25 communities across three states. Better welfare (specifically body condition and skin alteration scores) was seen in communities where a history of combined free veterinary treatment and educational initiatives had taken place compared to those that had only received veterinary treatment or control communities. The social transfer of welfare knowledge was also higher in these communities, suggesting that the discussion and transfer of equid welfare advice within communities can act as a mechanism to disseminate good welfare practices more widely. Our results suggest that using a combined approach may enhance the success of welfare initiatives, a finding that may impact future NGO programming.

Comparison of working equid welfare across three regions of Mexico.

BACKGROUND: Factors affecting working equid welfare are wide-ranging and reflect cultural, economic and climatic conditions, the type of work equids are used for, and individual differences in the practices of their handlers. In Mexico working equids are widely used for facilitating agricultural activities, however, welfare issues are common. OBJECTIVES: To assess working equids across three communities in Mexico, identify predominant welfare problems and document how these problems vary across locations, associated working roles and species type. STUDY DESIGN: Cross-sectional survey. METHODS: The study combined the administration of a wide-ranging questionnaire to equid handlers/owners and a welfare assessment of their animal. 120 equid owners were asked about their equid management practices, the working conditions and health status of their animal. The welfare of their equids (56 donkeys, 7 mules, 57 horses) was assessed by evaluating body condition, signs of illness or injury and behavioural indicators. RESULTS: Welfare varied by species, working role, sex and location. The poorest welfare was seen in one of the two arid regions (the third location having a tropical climate). Donkeys had poorer welfare than horses, and equids used for packing had poorer welfare than those used for riding and agroforestry. Overall poor body condition and wounds were the most common problems seen. MAIN LIMITATIONS: Work type, species type and location strongly co-varied, thus the impact of each factor could not be assessed in isolation. The sample size was relatively small. CONCLUSIONS: Results showed significant regional variations in welfare, suggesting that environmental and/or cultural variations are producing a major effect on welfare.

Slow Blink Eye Closure in Shelter Cats is Related to Quicker Adoption.

The process of domestication is likely to have led to the development of adaptive interspecific social abilities in animals. Such abilities are particularly interesting in less gregarious animals, such as cats. One notable social behaviour that cats exhibit in relation to humans is the slow blink sequence, which our previous research suggests can function as a form of positive communication between cats and humans. This behaviour involves the production of successive half blinks followed by either a prolonged narrowing of the eye or an eye closure. The present study investigates how cat (n = 18) slow blink sequences might affect human preferences during the adoption of shelter cats. Our study specifically tested (1) whether cats' propensity to respond to human-initiated slow blinking was associated with their speed of rehoming from a shelter environment, and (2) whether cats' anxiety around humans was related to their tendency to slow blink. Our experiments demonstrated that cats that showed an increased number of and longer eye closures in response to human slow blinks were rehomed faster, and that nervous cats, who had been identified as needing desensitisation to humans, tended to spend more time producing slow blink sequences in response to human slow blinks than a non-desensitisation group. Collectively, these results suggest that the cat slow blink sequence is perceived as positive by humans and may have a dual function-occurring in both affiliative and submissive contexts.

The role of cat eye narrowing movements in cat-human communication.

Domestic animals are sensitive to human cues that facilitate inter-specific communication, including cues to emotional state. The eyes are important in signalling emotions, with the act of narrowing the eyes appearing to be associated with positive emotional communication in a range of species. This study examines the communicatory significance of a widely reported cat behaviour that involves eye narrowing, referred to as the slow blink sequence. Slow blink sequences typically involve a series of half-blinks followed by either a prolonged eye narrow or an eye closure. Our first experiment revealed that cat half-blinks and eye narrowing occurred more frequently in response to owners' slow blink stimuli towards their cats (compared to no owner-cat interaction). In a second experiment, this time where an experimenter provided the slow blink stimulus, cats had a higher propensity to approach the experimenter after a slow blink interaction than when they had adopted a neutral expression. Collectively, our results suggest that slow blink sequences may function as a form of positive emotional communication between cats and humans.

Documenting the Welfare and Role of Working Equids in Rural Communities of Portugal and Spain.

Recently, the need for a more holistic approach to welfare assessment has been highlighted. This is particularly pertinent in the case of working equids who provide vital support for human livelihoods, often in low- to middle-income countries, yet suffer from globally low standards of welfare. This study aimed to provide insight into the welfare status and traditional use of working equids in rural Western European communities using the new EARS welfare tool, designed to provide a broad view of the welfare of working equids and the context in which they are found. Other questions on the topics of equid management practices, social transmission of expertise, environmental stressors, and traditions, alongside physical and behavioural welfare assessments were also included to explore the impact of these wide-ranging factors on an understudied population of working equids. The protocol was trialled on 60 working equid owners from communities in Portugal and Spain where, despite the decline in traditional agricultural practices and livestock keeping, donkeys and mules remain working animals. Many owners stated that the help donkeys provided was invaluable, and donkeys were considered to be important for both farming and daily life. However, participants also recognised that the traditional agricultural way of life was dying out, providing insights into the traditional practices, community structure, and beliefs of equid owners. Questions investigating the social networks and social transfer of information within the villages were effective in finding local sources of equid knowledge. Overall, welfare was deemed fair, and the protocol enabled the identification of the most prevalent welfare problems within the communities studied, in this case obesity and the use of harmful practices. The findings suggest that the new protocol was feasible and detail how contextual factors may influence equid welfare. Increasing understanding of the cultural context, social structure, and attitudes within a community, alongside more traditional investigations of working practices and animal management, may, in the future, help to make equid welfare initiatives more effective.

Domestic horses (Equus caballus) discriminate between negative and positive human nonverbal vocalisations.

The ability to discriminate between emotion in vocal signals is highly adaptive in social species. It may also be adaptive for domestic species to distinguish such signals in humans. Here we present a playback study investigating whether horses spontaneously respond in a functionally relevant way towards positive and negative emotion in human nonverbal vocalisations. We presented horses with positively- and negatively-valenced human vocalisations (laughter and growling, respectively) in the absence of all other emotional cues. Horses were found to adopt a freeze posture for significantly longer immediately after hearing negative versus positive human vocalisations, suggesting that negative voices promote vigilance behaviours and may therefore be perceived as more threatening. In support of this interpretation, horses held their ears forwards for longer and performed fewer ear movements in response to negative voices, which further suggest increased vigilance. In addition, horses showed a right-ear/left-hemisphere bias when attending to positive compared with negative voices, suggesting that horses perceive laughter as more positive than growling. These findings raise interesting questions about the potential for universal discrimination of vocal affect and the role of lifetime learning versus other factors in interspecific communication.

Animals Remember Previous Facial Expressions that Specific Humans Have Exhibited.

For humans, facial expressions are important social signals, and how we perceive specific individuals may be influenced by subtle emotional cues that they have given us in past encounters. A wide range of animal species are also capable of discriminating the emotions of others through facial expressions [1-5], and it is clear that remembering emotional experiences with specific individuals could have clear benefits for social bonding and aggression avoidance when these individuals are encountered again. Although there is evidence that non-human animals are capable of remembering the identity of individuals who have directly harmed them [6, 7], it is not known whether animals can form lasting memories of specific individuals simply by observing subtle emotional expressions that they exhibit on their faces. Here we conducted controlled experiments in which domestic horses were presented with a photograph of an angry or happy human face and several hours later saw the person who had given the expression in a neutral state. Short-term exposure to the facial expression was enough to generate clear differences in subsequent responses to that individual (but not to a different mismatched person), consistent with the past angry expression having been perceived negatively and the happy expression positively. Both humans were blind to the photograph that the horses had seen. Our results provide clear evidence that some non-human animals can effectively eavesdrop on the emotional state cues that humans reveal on a moment-to-moment basis, using their memory of these to guide future interactions with particular individuals.

Author Correction: Domestic horses (Equus caballus) prefer to approach humans displaying a submissive body posture rather than a dominant body posture.

In the original publication, data availability text was incorrectly published. The correct text should read as below.

Domestic horses (Equus caballus) prefer to approach humans displaying a submissive body posture rather than a dominant body posture.

Signals of dominance and submissiveness are central to conspecific communication in many species. For domestic animals, sensitivities to these signals in humans may also be beneficial. We presented domestic horses with a free choice between two unfamiliar humans, one adopting a submissive and the other a dominant body posture, with vocal and facial cues absent. Horses had previously been given food rewards by both human demonstrators, adopting neutral postures, to encourage approach behaviour. Across four counterbalanced test trials, horses showed a significant preference for approaching the submissive posture in both the first trial and across subsequent trials, and no individual subject showed an overall preference for dominant postures. There was no significant difference in latency to approach the two postures. This study provides novel evidence that domestic horses may spontaneously discriminate between, and attribute communicative significance to, human body postures of dominance; and further, that familiarity with the signaller is not a requirement for this response. These findings raise interesting questions about the plasticity of social signal perception across the species barrier.

Functionally relevant responses to human facial expressions of emotion in the domestic horse (Equus caballus).

Whether non-human animals can recognize human signals, including emotions, has both scientific and applied importance, and is particularly relevant for domesticated species. This study presents the first evidence of horses' abilities to spontaneously discriminate between positive (happy) and negative (angry) human facial expressions in photographs. Our results showed that the angry faces induced responses indicative of a functional understanding of the stimuli: horses displayed a left-gaze bias (a lateralization generally associated with stimuli perceived as negative) and a quicker increase in heart rate (HR) towards these photographs. Such lateralized responses towards human emotion have previously only been documented in dogs, and effects of facial expressions on HR have not been shown in any heterospecific studies. Alongside the insights that these findings provide into interspecific communication, they raise interesting questions about the generality and adaptiveness of emotional expression and perception across species.

The Responses of Young Domestic Horses to Human-Given Cues.

It has been suggested that the process of domestication, at least in some species, has led to an innate predisposition to be skilled at reading human communicative and attentional cues. Adult domestic horses (Equus caballus) are highly sensitive to subtle bodily cues when determining if a person is attending to them but they are less adept at using human communicative cues in object choice tasks. Here we provide the first study into the ontogeny of such skills in order to gain insights into the mechanisms underlying these abilities. Compared with adult horses, youngsters under the age of three could use body orientation but not more subtle cues such as head movement and open/closed eyes to correctly choose an attentive person to approach for food. Across two object choice experiments, the performance of young horses was comparable to that of adult horses - subjects were able to correctly choose a rewarded bucket using marker placement, pointing and touching cues but could not use body orientation, gaze, elbow pointing or tapping cues. Taken together these results do not support the theory that horses possess an innate predisposition to be particularly skilled at using human cues. Horses' ability to determine whether humans are attending to them using subtle body cues appears to require significant experience to fully develop and their perhaps less remarkable use of limited cues in object choice tasks, although present at a much earlier age, is likely to reflect a more general learning ability related to stimulus enhancement rather than a specific 'human-reading' skill.

Spatial cognition and perseveration by horses, donkeys and mules in a simple A-not-B detour task.

We investigated perseveration and detour behaviour in 36 equids (Equus caballus, E. asinus, E. caballus × E. asinus) and compared these data to those of a previous study on domestic dogs (Canis familiaris). The animals were required to make a detour through a gap at one end of a straight barrier in order to reach a visible target. After one, two, three or four repeats (A trials), the gap was moved to the opposite end of the barrier (B trials). We recorded initial deviations from the correct solution path and the latency to crossing the barrier. In the A trials, mules crossed the barrier significantly faster than their parental species, the horses and donkeys. In the B trials, following the change of gap location, all species showed a reduction in performance. Both dogs and horses exhibited significant spatial perseveration, going initially to the previous gap location. Donkeys and mules, however, performed at chance level. Our results suggest that hybrid vigour in mules extends to spatial abilities.

Social relations in a mixed group of mules, ponies and donkeys reflect differences in equid type.

Donkeys and mules are frequently kept as companion animals for horses and ponies, with these different equids often being considered a homogenous group. However, the extent to which domestic equids form inter-specific bonds and display similar social behaviour when living in a mixed herd has not previously been studied. Here we compare the social organization of these three (sub)species when housed together, providing the first systematic analysis of how genetic hybridization is expressed in the social behaviour of mules. A group of 16 mules, donkeys and ponies was observed for 70h and preferred associates, dominance rank and the linearity of the group's hierarchy was determined. The different equids formed distinct affiliative groups that were ordered in a linear hierarchy with ponies as the most dominant, mules in the middle ranks and donkeys in the lowest ranks. Within each equid subgroup, the strength of the hierarchy also varied. Thus in the present study, the three (sub)species displayed different social organization and levels of dominance and preferred to associate with animals of the same equid type, given the opportunity. These results suggest that different domestic equid (sub)species display variations in social behaviour that are likely to have a strong genetic basis.

Cross-modal individual recognition in domestic horses (Equus caballus) extends to familiar humans.

It has recently been shown that some non-human animals can cross-modally recognize members of their own taxon. What is unclear is just how plastic this recognition system can be. In this study, we investigate whether an animal, the domestic horse, is capable of spontaneous cross-modal recognition of individuals from a morphologically very different species. We also provide the first insights into how cross-modal identity information is processed by examining whether there are hemispheric biases in this important social skill. In our preferential looking paradigm, subjects were presented with two people and playbacks of their voices to determine whether they were able to match the voice with the person. When presented with familiar handlers subjects could match the specific familiar person with the correct familiar voice. Horses were significantly better at performing the matching task when the congruent person was standing on their right, indicating marked hemispheric specialization (left hemisphere bias) in this ability. These results are the first to demonstrate that cross-modal recognition in animals can extend to individuals from phylogenetically very distant species. They also indicate that processes governed by the left hemisphere are central to the cross-modal matching of visual and auditory information from familiar individuals in a naturalistic setting.