"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.

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.

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.

"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.

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.

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.

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.

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.

Dogs (Canis familiaris) stick to what they have learned rather than conform to their conspecifics' behavior.

In recent years, an increasing number of studies has investigated majority influence in nonhuman animals. However, due to both terminological and methodological issues, evidence for conformity in nonhuman animals is scarce and controversial. Preliminary evidence suggests that wild birds, wild monkeys, and fish show conformity, that is, forgoing personal information in order to copy the majority. By contrast, chimpanzees seem to lack this tendency. The present study is the first to examine whether dogs (Canis familiaris) show conformity. Specifically, we tested whether dogs conform to a majority of conspecifics rather than stick to what they have previously learned. After dogs had acquired a behavioral preference via training (i.e., shaping), they were confronted with counter-preferential behavior of either no, one or three conspecifics. Traditional frequentist analyses show that the dogs' behavior did not differ significantly between the three conditions. Complementary Bayesian analyses suggest that our data provide moderate evidence for the null hypothesis. In conclusion, our results suggest that dogs stick to what they have learned rather than conform to the counter-preferential behavior of others. We discuss the possible statistical and methodological limitations of this finding. Furthermore, we take a functional perspective on conformity and discuss under which circumstances dogs might show conformity after all.

Human attention affects facial expressions in domestic dogs.

Most mammalian species produce facial expressions. Historically, animal facial expressions have been considered inflexible and involuntary displays of emotional states rather than active attempts to communicate with others. In the current study, we aimed to test whether domestic dog facial expressions are subject to audience effects and/ or changes in response to an arousing stimulus (e.g. food) alone. We presented dogs with an experimental situation in which a human demonstrator was either attending to them or turned away, and varied whether she presented food or not. Dogs produced significantly more facial movements when the human was attentive than when she was not. The food, however, as a non-social but arousing stimulus, did not affect the dogs' behaviour. The current study is therefore evidence that dogs are sensitive to the human's attentional state when producing facial expressions, suggesting that facial expressions are not just inflexible and involuntary displays of emotional states, but rather potentially active attempts to communicate with others.

The effects of domestication and ontogeny on cognition in dogs and wolves.

Cognition is one of the most flexible tools enabling adaptation to environmental variation. Living close to humans is thought to influence social as well as physical cognition of animals throughout domestication and ontogeny. Here, we investigated to what extent physical cognition and two domains of social cognition of dogs have been affected by domestication and ontogeny. To address the effects of domestication, we compared captive wolves (n = 12) and dogs (n = 14) living in packs under the same conditions. To explore developmental effects, we compared these dogs to pet dogs (n = 12) living in human families. The animals were faced with a series of object-choice tasks, in which their response to communicative, behavioural and causal cues was tested. We observed that wolves outperformed dogs in their ability to follow causal cues, suggesting that domestication altered specific skills relating to this domain, whereas developmental effects had surprisingly no influence. All three groups performed similarly in the communicative and behavioural conditions, suggesting higher ontogenetic flexibility in the two social domains. These differences across cognitive domains need to be further investigated, by comparing domestic and non-domesticated animals living in varying conditions.

Prelinguistic human infants and great apes show different communicative strategies in a triadic request situation.

In the present research, we investigate the communicative strategies of 20 month old human infants and great apes when requesting rewards from a human experimenter. Infants and apes both adapted their signals to the attentional state of the experimenter as well as to the location of the reward. Yet, while infants frequently positioned themselves in front of the experimenter and pointed towards a distant reward, apes either remained in the experimenter's line of sight and pointed towards him or moved out of sight and pointed towards the reward. Further, when pointing towards a reward that was placed at a distance from the experimenter, only the infants, and not the apes, took the experimenter's attentional state into account. These results demonstrate that prelinguistic human infants and nonhuman apes use different means when guiding others' attention to a location; indicating that differing cognitive mechanisms may underlie their pointing gestures.

Do Dogs Provide Information Helpfully?

Dogs are particularly skilful during communicative interactions with humans. Dogs' abilities to use human communicative cues in cooperative contexts outcompete those of other species, and might be the result of selection pressures during domestication. Dogs also produce signals to direct the attention of humans towards outside entities, a behaviour often referred to as showing behaviour. This showing behaviour in dogs is thought to be something dogs use intentionally and referentially. However, there is currently no evidence that dogs communicate helpfully, i.e. to inform an ignorant human about a target that is of interest to the human but not to the dog. Communicating with a helpful motive is particularly interesting because it might suggest that dogs understand the human's goals and need for information. In study 1, we assessed whether dogs would abandon an object that they find interesting in favour of an object useful for their human partner, a random novel distractor, or an empty container. Results showed that it was mainly self-interest that was driving the dogs' behaviour. The dogs mainly directed their behaviour towards the object they had an interest in, but dogs were more persistent when showing the object relevant to the human, suggesting that to some extent they took the humans interest into account. Another possibility is that dogs' behaviour was driven by an egocentric motivation to interact with novel targets and that the dogs' neophila might have masked their helpful tendencies. Therefore, in study 2 the dogs had initial access to both objects, and were expected to indicate only one (relevant or distractor). The human partner interacted with the dog using vocal communication in half of the trials, and remaining silent in the other half. Dogs from both experimental groups, i.e. indicating the relevant object or indicating the distractor, established joint attention with the human. However, the human's vocal communication and the presence of the object relevant to the human increased the persistency of showing, supporting the hypothesis that the dogs understood the objects' relevance to the human. We propose two non-exclusive explanations. These results might suggest that informative motives could possibly underlie dogs' showing. It is also possible that dogs might have indicated the location of the hidden object because they recognised it as the target of the human's search. This would be consistent with taking into account the objects' relevance, without necessarily implying that the dogs understood the human's state of knowledge.

Citizen Science as a New Tool in Dog Cognition Research.

Family dogs and dog owners offer a potentially powerful way to conduct citizen science to answer questions about animal behavior that are difficult to answer with more conventional approaches. Here we evaluate the quality of the first data on dog cognition collected by citizen scientists using the Dognition.com website. We conducted analyses to understand if data generated by over 500 citizen scientists replicates internally and in comparison to previously published findings. Half of participants participated for free while the other half paid for access. The website provided each participant a temperament questionnaire and instructions on how to conduct a series of ten cognitive tests. Participation required internet access, a dog and some common household items. Participants could record their responses on any PC, tablet or smartphone from anywhere in the world and data were retained on servers. Results from citizen scientists and their dogs replicated a number of previously described phenomena from conventional lab-based research. There was little evidence that citizen scientists manipulated their results. To illustrate the potential uses of relatively large samples of citizen science data, we then used factor analysis to examine individual differences across the cognitive tasks. The data were best explained by multiple factors in support of the hypothesis that nonhumans, including dogs, can evolve multiple cognitive domains that vary independently. This analysis suggests that in the future, citizen scientists will generate useful datasets that test hypotheses and answer questions as a complement to conventional laboratory techniques used to study dog psychology.

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.

Generalize or personalize--do dogs transfer an acquired rule to novel situations and persons?

Recent studies have raised the question of whether dogs, like human infants, comprehend an established rule as generalizable, normative knowledge or rather as episodic information, existing only in the immediate situation. In the current study we tested whether dogs disobeyed a prohibition to take a treat (i) in the presence of the communicator of the ban, (ii) after a temporary absence of the communicator, and (iii) in the presence of a novel person. Dogs disobeyed the rule significantly more often when the communicator left the room for a moment or when they were faced with a new person, than when she stayed present in the room. These results indicate that dogs "forget" a rule as soon as the immediate human context becomes disrupted.