How wolves were first domesticated is unknown. One hypothesis suggests that wolves underwent a process of self-domestication by tolerating human presence and taking advantage of scavenging possibilities. The puppy-like physical and behavioural traits seen in dogs are thought to have evolved later, as a byproduct of selection against aggression. Using speed of selection from rehoming shelters as a proxy for artificial selection, we tested whether paedomorphic features give dogs a selective advantage in their current environment. Dogs who exhibited facial expressions that enhance their neonatal appearance were preferentially selected by humans. Thus, early domestication of wolves may have occurred not only as wolf populations became tamer, but also as they exploited human preferences for paedomorphic characteristics. These findings, therefore, add to our understanding of early dog domestication as a complex co-evolutionary process.

The commonality of facial expressions of emotion has been studied in different species since Darwin, with most of the research focusing on closely related primate species. However, it is unclear to what extent there exists common facial expression in species more phylogenetically distant, but sharing a need for common interspecific emotional understanding. Here we used the objective, anatomically-based tools, FACS and DogFACS (Facial Action Coding Systems), to quantify and compare human and domestic dog facial expressions in response to emotionally-competent stimuli associated with different categories of emotional arousal. We sought to answer two questions: Firstly, do dogs display specific discriminatory facial movements in response to different categories of emotional stimuli? Secondly, do dogs display similar facial movements to humans when reacting in emotionally comparable contexts? We found that dogs displayed distinctive facial actions depending on the category of stimuli. However, dogs produced different facial movements to humans in comparable states of emotional arousal. These results refute the commonality of emotional expression across mammals, since dogs do not display human-like facial expressions. Given the unique interspecific relationship between dogs and humans, two highly social but evolutionarily distant species sharing a common environment, these findings give new insight into the origin of emotion expression.

The domestic cat (Felis silvestris catus) is quickly becoming the most popular animal companion in the world. The evolutionary processes that occur during domestication are known to have wide effects on the morphology, behaviour, cognition and communicative abilities of a species. Since facial expression is central to human communication, it is possible that cat facial expression has been subject to selection during domestication. Standardised measurement techniques to study cat facial expression are, however, currently lacking. Here, as a first step to enable cat facial expression to be studied in an anatomically based and objective way, CatFACS (Cat Facial Action Coding System) was developed. Fifteen individual facial movements (Action Units), six miscellaneous movements (Action Descriptors) and seven Ear Action Descriptors were identified in the domestic cat. CatFACS was then applied to investigate the impact of cat facial expression on human preferences in an adoption shelter setting. Rehoming speed from cat shelters was used as a proxy for human selective pressure. The behaviour of 106 cats ready for adoption in three different shelters was recorded during a standardised encounter with an experimenter. This experimental setup aimed to mimic the first encounter of a cat with a potential adopter, i.e. an unfamiliar human. Each video was coded for proximity to the experimenter, body movements, tail movements and face movements. Cat facial movements were not related to rehoming speed, suggesting that cat facial expression may not have undergone significant selection. In contrast, rubbing frequency was positively related to rehoming speed. The findings suggest that humans are more influenced by overt prosocial behaviours than subtle facial expression in domestic cats.

Comparing homologous expressions between species can shed light on the phylogenetic and functional changes that have taken place during evolution. To assess homology across species we must approach primate facial expressions in an anatomical, systematic, and standardized way. The Facial Action Coding System (FACS), a widely used muscle-based tool for analyzing human facial expressions, has recently been adapted for chimpanzees (Pan troglodytes: ChimpFACS), rhesus macaques (Macaca mulatta: MaqFACS), and gibbons (GibbonFACS). Here, we present OrangFACS, a FACS adapted for orangutans (Pongo spp.). Orangutans are the most arboreal and the least social great ape, so their visual communication has been assumed to be less important than vocal communication and is little studied. We scrutinized the facial anatomy of orangutans and coded videos of spontaneous orangutan behavior to identify independent movements: Action Units (AUs) and Action Descriptors (ADs). We then compared these facial movements with movements of homologous muscles in humans, chimpanzees, macaques, and gibbons. We also noted differences related to sexual dimorphism and developmental stages in orangutan facial morphology. Our results

Primate facial expressions are widely accepted as underpinned by reflexive emotional processes and not under voluntary control. In contrast, other modes of primate communication, especially gestures, are widely accepted as underpinned by intentional, goal-driven cognitive processes. One reason for this distinction is that production of primate gestures is often sensitive to the attentional state of the recipient, a phenomenon used as one of the key behavioural criteria for identifying intentionality in signal production. The reasoning is that modifying/producing a signal when a potential recipient is looking could demonstrate that the sender intends to communicate with them. Here, we show that the production of a primate facial expression can also be sensitive to the attention of the play partner. Using the orangutan (Pongo pygmaeus) Facial Action Coding System (OrangFACS), we demonstrate that facial movements are more intense and more complex when recipient attention is directed towards the sender. Therefore, production of the playface is not an automated response to play (or simply a play behaviour itself) and is instead produced flexibly depending on the context. If sensitivity to attentional stance is a good indicator of intentionality, we must also conclude that the orangutan playface is intentionally produced. However, a number of alternative, lower level interpretations for flexible production of signals in response to the attention of another are discussed. As intentionality is a key feature of human language, claims of intentional communication in related primate species are powerful drivers in language evolution debates, and thus caution in identifying intentionality is important.

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