Social gaze has received much attention in social cognition research in both human and non-human animals. Gaze following appears to be a central skill for acquiring social information, such as the location of food and predators, but can also draw attention to important social interactions, which in turn promotes the evolution of more complex socio-cognitive processes such as theory of mind and social learning. In the past decades, a large number of studies has been conducted in this field introducing differing methodologies. Thereby, various factors influencing the results of gaze following experiments have been identified. This review provides an overview of the advances in the study of gaze following, but also highlights some limitations within the research area. The majority of gaze following studies on animals have focused on primates and canids, which limits evolutionary interpretations to only a few and closely related evolutionary lineages. This review incorporates new insights gained from previously understudied taxa, such as fishes, reptiles, and birds, but it will also provide a brief outline of mammal studies. We propose that the foundations of gaze following emerged early in evolutionary history. Basic, reflexive co-orienting responses might have already evolved in fishes, which would explain the ubiquity of gaze following seen in the amniotes. More complex skills, such as geometrical gaze following and the ability to form social predictions based on gaze, seem to have evolved separately at least two times and appear to be correlated with growing complexity in brain anatomy such as increased numbers of brain neurons. However, more studies on different taxa in key phylogenetic positions are needed to better understand the evolutionary history of this fundamental socio-cognitive skill.

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Preening behaviours are widespread in extant birds. While most birds appear to autopreen (self-directed preening), allopreening (preening directed at conspecifics) seems to have emerged only in certain species, but across many families. Allopreening has been hypothesised to reinforce mutual relationships and cooperation between individuals, and to underpin various socio-cognitive abilities. Palaeognathae is a bird group exhibiting neurocognitively plesiomorphic traits compared to other birds. They share many features with non-avian paravian dinosaurs and are thus important for the study of cognitive evolution in birds. Despite this, and the important correlation of allopreening with many complicated social behaviours, allopreening has not been systematically studied in Palaeognathae. Therefore, we examined the preening behaviours in four species of palaeognaths: common ostriches (Struthio camelus), greater rheas (Rhea americana), emus (Dromaius novaehollandiae), and elegant crested tinamous (Eudromia elegans). We compared findings with common ravens (Corvus corax), a neognath species known for its allopreening and complex social cognition. We found autopreening, but no allopreening, in the palaeognath species, while both autopreening and allopreening was found in common ravens. The absence of allopreening in Palaeognathae suggests an emergence of this behaviour within Neognathae. We contextualise our results in relation to the socio-cognitive underpinnings of allopreening and its implications for the understanding of the evolution of socio-cognitive abilities in non-avian paravian dinosaurs and early birds.

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