Perception of optical illusions in ungulates: insights from goats, sheep, guanacos and llamas.

Optical illusions have long been used in behavioural studies to investigate the perceptual mechanisms underlying vision in animals. So far, three studies have focused on ungulates, providing evidence that they may be susceptible to some optical illusions, in a way similar to humans. Here, we used two food-choice tasks to study susceptibility to the Müller-Lyer and Delboeuf illusions in 17 captive individuals belonging to four ungulate species (Lama guanicoe, Lama glama, Ovis aries, Capra hircus). At the group level, there was a significant preference for the longer/larger food over the shorter/smaller one in control trials. Additionally, the whole group significantly preferred the food stick between two inward arrowheads over an identical one between two outward arrowheads in experimental trials of the Müller-Lyer task, and also preferred the food on the smaller circle over an identical one on the larger circle in the experimental trials of the Delboeuf task. Group-level analyses further showed no significant differences across species, although at the individual level we found significant variation in performance. Our findings suggest that, in line with our predictions, ungulates are overall susceptible to the Müller-Lyer and the Delboeuf illusions, and indicate that the perceptual mechanisms underlying size estimation in artiodactyls might be similar to those of other species, including humans.

Evidence of object permanence, short-term spatial memory, causality, understanding of object properties and gravity across five different ungulate species.

In their natural environment, animals face a variety of ecological and social challenges, which might be linked to the emergence of different cognitive skills. To assess inter-specific variation in cognitive skills, we used ungulates as a study model, testing a total of 26 captive individuals across 5 different species (i.e., dwarf goats, Capra aegagrus hircus, llamas, Lama glama, guanacos, Lama guanicoe, zebras, Equus grevyi, and rhinos, Diceros bicornis michaeli). Across species, we used the same well-established experimental procedures to test individuals' performance in naïve physics tasks, i.e. object permanence, short-term spatial memory, causality, understanding of object properties, and gravity. Our results revealed that study subjects showed object permanence, were able to remember the position of hidden food after up to 60 s, and inferred the position of hidden food from the sound produced or not produced when shaking containers. Moreover, they showed an understanding of basic object properties, being able to locate objects hidden behind occluders based on their size and inclination, and could reliably follow the trajectory of falling objects across different conditions. Finally, inter-specific differences were limited to the understanding of object properties, and suggest that domesticated species as goats might perform better than non-domesticated ones in tasks requiring these skills. These results provide new information on the cognitive skills of a still understudied taxon and confirm ungulates as a promising taxon for the comparative study of cognitive evolution.