"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.

Evolution of facial muscle anatomy in dogs.

Domestication shaped wolves into dogs and transformed both their behavior and their anatomy. Here we show that, in only 33,000 y, domestication transformed the facial muscle anatomy of dogs specifically for facial communication with humans. Based on dissections of dog and wolf heads, we show that the levator anguli oculi medialis, a muscle responsible for raising the inner eyebrow intensely, is uniformly present in dogs but not in wolves. Behavioral data, collected from dogs and wolves, show that dogs produce the eyebrow movement significantly more often and with higher intensity than wolves do, with highest-intensity movements produced exclusively by dogs. Interestingly, this movement increases paedomorphism and resembles an expression that humans produce when sad, so its production in dogs may trigger a nurturing response in humans. We hypothesize that dogs with expressive eyebrows had a selection advantage and that "puppy dog eyes" are the result of selection based on humans' preferences.

Age influences domestic dog cognitive performance independent of average breed lifespan.

Across mammals, increased body size is positively associated with lifespan. However, within species, this relationship is inverted. This is well illustrated in dogs (Canis familiaris), where larger dogs exhibit accelerated life trajectories: growing faster and dying younger than smaller dogs. Similarly, some age-associated traits (e.g., growth rate and physiological pace of aging) exhibit accelerated trajectories in larger breeds. Yet, it is unknown whether cognitive performance also demonstrates an accelerated life course trajectory in larger dogs. Here, we measured cognitive development and aging in a cross-sectional study of over 4000 dogs from 66 breeds using nine memory and decision-making tasks performed by citizen scientists as part of the Dognition project. Specifically, we tested whether cognitive traits follow a compressed (accelerated) trajectory in larger dogs, or the same trajectory for all breeds, which would result in limited cognitive decline in larger breeds. We found that all breeds, regardless of size or lifespan, tended to follow the same quadratic trajectory of cognitive aging-with a period of cognitive development in early life and decline in later life. Taken together, our results suggest that cognitive performance follows similar age-related trajectories across dog breeds, despite remarkable variation in developmental rates and lifespan.

Absolute brain size predicts dog breed differences in executive function.

Large-scale phylogenetic studies of animal cognition have revealed robust links between absolute brain volume and species differences in executive function. However, past comparative samples have been composed largely of primates, which are characterized by evolutionarily derived neural scaling rules. Therefore, it is currently unknown whether positive associations between brain volume and executive function reflect a broad-scale evolutionary phenomenon, or alternatively, a unique consequence of primate brain evolution. Domestic dogs provide a powerful opportunity for investigating this question due to their close genetic relatedness, but vast intraspecific variation. Using citizen science data on more than 7000 purebred dogs from 74 breeds, and controlling for genetic relatedness between breeds, we identify strong relationships between estimated absolute brain weight and breed differences in cognition. Specifically, larger-brained breeds performed significantly better on measures of short-term memory and self-control. However, the relationships between estimated brain weight and other cognitive measures varied widely, supporting domain-specific accounts of cognitive evolution. Our results suggest that evolutionary increases in brain size are positively associated with taxonomic differences in executive function, even in the absence of primate-like neuroanatomy. These findings also suggest that variation between dog breeds may present a powerful model for investigating correlated changes in neuroanatomy and cognition among closely related taxa.

Two-year-old children but not domestic dogs understand communicative intentions without language, gestures, or gaze.

Infants can see someone pointing to one of two buckets and infer that the toy they are seeking is hidden inside. Great apes do not succeed in this task, but, surprisingly, domestic dogs do. However, whether children and dogs understand these communicative acts in the same way is not yet known. To test this possibility, an experimenter did not point, look, or extend any part of her body towards either bucket, but instead lifted and shook one via a centrally pulled rope. She did this either intentionally or accidentally, and did or did not address her act to the subject using ostensive cues. Young 2-year-old children but not dogs understood the experimenter's act in intentional conditions. While ostensive pulling of the rope made no difference to children's success, it actually hindered dogs' performance. We conclude that while human children may be capable of inferring communicative intent from a wide variety actions, so long as these actions are performed intentionally, dogs are likely to be less flexible in this respect. Their understanding of communicative intention may be more dependent upon bodily markers of communicative intent, including gaze, orientation, extended limbs, and vocalizations. This may be because humans have come under selective pressure to develop skills for communicating with absent interlocutors - where bodily co-presence is not possible.

Understanding others' preferences: A comparison across primate species and human societies.

We investigated children's and non-human great apes' ability to anticipate others' choices from their evident food preferences-regardless of whether these preferences deviate or align with one's own. We assessed children from three culturally-diverse societies (Namibia, Germany, and Samoa; N = 71; age range = 5-11) and four non-human great ape species (chimpanzees (Pan troglodytes), bonobos (Pan paniscus), gorillas (Gorilla gorilla), and orangutans (Pongo abelii); N = 25; age range = 7-29) regarding their choices in a dyadic food-retrieval task. Across conditions, participants' preferences were either aligned (same preference condition) or opposed (opposite preference condition) to those of their competitors. Children across societies altered their choices based on their competitor's preferences, indicating a cross-culturally recurrent capacity to anticipate others' choices relying on preferences-based inferences. In contrast to human children, all non-human great apes chose according to their own preferences but independent of those of their competitors. In sum, these results suggest that the tendency to anticipate others' choices based on their food preferences is cross-culturally robust and, among the great apes, most likely specific to humans.

Dogs steal in the dark.

All current evidence of visual perspective taking in dogs can possibly be explained by dogs reacting to certain stimuli rather than understanding what others see. In the current study, we set up a situation in which contextual information and social cues are in conflict. A human always forbade the dog from taking a piece of food. The part of the room being illuminated was then varied, for example, either the area where the human was seated or the area where the food was located was lit. Results show that dogs steal significantly more food when it is dark compared to when it is light. While stealing forbidden food the dog's behaviour also depends on the type of illumination in the room. Illumination around the food, but not the human, affected the dogs' behaviour. This indicates that dogs do not take the sight of the human as a signal to avoid the food. It also cannot be explained by a low-level associative rule of avoiding illuminated food which dogs actually approach faster when they are in private. The current finding therefore raises the possibility that dogs take into account the human's visual access to the food while making their decision to steal it.

Do domestic dogs interpret pointing as a command?

Domestic dogs comprehend human gestural communication flexibly, particularly the pointing gesture. Here, we examine whether dogs interpret pointing informatively, that is, as simply providing information, or rather as a command, for example, ordering them to move to a particular location. In the first study a human pointed toward an empty cup. In one manipulation, the dog either knew or did not know that the designated cup was empty (and that the other cup actually contained the food). In another manipulation, the human (as authority) either did or did not remain in the room after pointing. Dogs ignored the human's gesture if they had better information, irrespective of the authority's presence. In the second study, we varied the level of authority of the person pointing. Sometimes this person was an adult, and sometimes a young child. Dogs followed children's pointing just as frequently as they followed adults' pointing (and ignored the dishonest pointing of both), suggesting that the level of authority did not affect their behavior. Taken together these studies suggest that dogs do not see pointing as an imperative command ordering them to a particular location. It is still not totally clear, however, if they interpret it as informative or in some other way.

Domestic dogs conceal auditory but not visual information from others.

A number of studies have shown that dogs are sensitive to a human's perspective, but it remains unclear whether they use an egocentric strategy to assess what humans perceive. We investigated whether dogs know what a human can see and hear, even when the dogs themselves are unable to see the human. Dogs faced a task in which forbidden food was placed in a tunnel that they could retrieve by using their paw. Whereas the dogs could not see the experimenter during their food retrieval attempts, the experimenter could potentially see the dog's paw. In the first experiment, dogs could choose between an opaque and a transparent side of the tunnel, and in the second experiment, they could choose between a silent and a noisy approach to the tunnel. The results showed that dogs preferred a silent approach to forbidden food but they did not hide their approach when they could not see a human present. We conclude that dogs probably rely on what they themselves can perceive when they assess what the human can see and hear.

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.

Gibbons (Hylobates pileatus, H. moloch, H. lar, Symphalangus syndactylus) follow human gaze, but do not take the visual perspective of others.

We investigated four gibbon species of two different genera (Hylobates pileatus, H. moloch, H. lar, Symphalangus syndactylus) in terms of their looking behavior in response to a human who either looked up or looked at the gibbon. Comparing those two conditions, gibbons as a group looked up more when the human was looking up, but they also performed more looks in other directions and thus generally looked more in this condition. Unlike great apes, gibbons did not respond differently between conditions when only the first look on every trial was considered. Furthermore, they did not perform double looks up to check where the human was looking and also did not habituate to the human's looks up. This suggests that gibbons co-orient with human gaze, but unlike great apes, they do not take the visual perspective of others.

How two word-trained dogs integrate pointing and naming.

Two word-trained dogs were presented with acts of reference in which a human pointed, named objects, or simultaneously did both. The question was whether these dogs would assume co-reference of pointing and naming and thus pick the pointed-to object. Results show that the dogs did indeed assume co-reference of pointing and naming in order to determine the reference of a spoken word, but they did so only when pointing was not in conflict with their previous word knowledge. When pointing and a spoken word conflicted, the dogs preferentially fetched the object by name. This is not surprising since they are trained to fetch objects by name. However, interestingly, in these conflict conditions, the dogs fetched the named objects only after they had initially approached the pointed-to object. We suggest that this shows that the word-trained dogs interpret pointing as a spatial directive, which they integrate into the fetching game, presumably assuming that pointing is relevant to finding the requested object.

How dogs know when communication is intended for them.

Domestic dogs comprehend human gestural communication in a way that other animal species do not. But little is known about the specific cues they use to determine when human communication is intended for them. In a series of four studies, we confronted both adult dogs and young dog puppies with object choice tasks in which a human indicated one of two opaque cups by either pointing to it or gazing at it. We varied whether the communicator made eye contact with the dog in association with the gesture (or whether her back was turned or her eyes were directed at another recipient) and whether the communicator called the dog's name (or the name of another recipient). Results demonstrated the importance of eye contact in human-dog communication, and, to a lesser extent, the calling of the dog's name--with no difference between adult dogs and young puppies--which are precisely the communicative cues used by human infants for identifying communicative intent. Unlike human children, however, dogs did not seem to comprehend the human's communicative gesture when it was directed to another human, perhaps because dogs view all human communicative acts as directives for the recipient.

"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.

Focus on the essential: all great apes know when others are being attentive.

When begging for food, all great ape species are sensitive to a human's attention. However, studies investigating which cues are relevant for chimpanzees to assess the attentional state of others have produced highly inconsistent results. Some have suggested chimpanzees differentiate attention based on the status of the face or even the eyes, while others have indicated that body posture alone is the relevant cue. Kaminski et al. (Anim Cogn 7:216-223, 2004) compared the behaviour of chimpanzees, bonobos and orangutans while begging for food from a human experimenter who systematically varied his face and body orientation. Their results indicated that both factors, face and body orientation, affect apes' begging behaviour. The authors claimed that while body orientation provides information about the experimenter's general disposition to offer food, the visibility of the face provides information about the human's attentional state. In the current study, we tested this hypothesis with all four great apes species. However, unlike Kaminski et al. (Anim Cogn 7:216-223, 2004), the experimenter was able to hand over food regardless of body orientation. The results show that as soon as the offering of the food was no longer restricted, the orientation of the face became the key factor. Therefore, we present the first evidence that all great ape species are able to assess the attentional state of a recipient based on the orientation of the face.

Different social motives in the gestural communication of chimpanzees and human children.

Both chimpanzees and human infants use the pointing gesture with human adults, but it is not clear if they are doing so for the same social motives. In two studies, we presented chimpanzees and human 25-month-olds with the opportunity to point for a hidden tool (in the presence of a non-functional distractor). In one condition it was clear that the tool would be used to retrieve a reward for the pointing subject (so the pointing was selfish or 'for-me'), whereas in the other condition it was clear that the tool would be used to retrieve the reward for the experimenter (so the pointing was helpful or 'for-you'). The chimpanzees pointed reliably only when they themselves benefited, whereas the human children pointed reliably no matter who benefited. These results are interpreted as evidence for the especially cooperative nature of human communication.

Domestic dogs: Born human whisperers.

Domestic dogs show socio-cognitive abilities that are not found in other species, including dogs' closest living relative, the wolf. From early on dogs are good at reading human gestures and pay attention to humans' eyes and voice. Two new studies suggest that a heritable component to dogs' social cognitive skills exists and that selection during domestication must have played a role in enhancing dogs' social cognitive skills.

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.

Breed Differences in Dog Cognition Associated with Brain-Expressed Genes and Neurological Functions.

Given their remarkable phenotypic diversity, dogs present a unique opportunity for investigating the genetic bases of cognitive and behavioral traits. Our previous work demonstrated that genetic relatedness among breeds accounts for a substantial portion of variation in dog cognition. Here, we investigated the genetic architecture of breed differences in cognition, seeking to identify genes that contribute to variation in cognitive phenotypes. To do so, we combined cognitive data from the citizen science project Dognition.com with published breed-average genetic polymorphism data, resulting in a dataset of 1654 individuals with cognitive phenotypes representing 49 breeds. We conducted a breed-average genome-wide association study to identify specific polymorphisms associated with breed differences in inhibitory control, communication, memory, and physical reasoning. We found five single nucleotide polymorphisms (SNPs) that reached genome-wide significance after Bonferroni correction, located in EML1, OR52E2, HS3ST5, a U6 spliceosomal RNA, and a long noncoding RNA. When we combined results across multiple SNPs within the same gene, we identified 188 genes implicated in breed differences in cognition. This gene set included more genes than expected by chance that were (1) differentially expressed in brain tissue and (2) involved in nervous system functions including peripheral nervous system development, Wnt signaling, presynapse assembly, and synaptic vesicle exocytosis. These results advance our understanding of the genetic underpinnings of complex cognitive phenotypes and identify specific genetic variants for further research.