Tortoises develop and overcome position biases in a reversal learning task

The capability of animals to alter their behaviour in response to novel or familiar stimuli, or behavioural flexibility, is strongly associated with their ability to learn in novel environments. Reptiles are capable of learning complex tasks and offer a unique opportunity to study the relationship between visual proficiency and behavioural flexibility. The focus of this study was to investigate the behavioural flexibility of red-footed tortoises and their ability to perform reversal learning. Reversal learning involves learning a particular discrimination task, after which the previously rewarded cue is reversed and then subjects perform the task with new reward contingencies. Red-footed tortoises were required to learn to recognise and approach visual cues within a Y-maze. Once subjects learned the visual discrimination, tortoises were required to successfully learn four reversals. Tortoises required significantly more trials to reach criterion (80% correct) in the first reversal, indicating the difficulty of unlearning the positive stimulus presented during training. Nevertheless, subsequent reversals required a similar number of sessions to the training stage, demonstrating that reversal learning improved up to a point. All subjects tested developed a position bias within the Y-maze that was absent prior to training, but most were able to exhibit reversal learning. Red-footed tortoises primarily adopted a win-stay choice strategy while learning the discrimination without much evidence for a lose-shift choice strategy, which may explain limits to their behavioural flexibility. However, improving performance across reversals while simultaneously overcoming a position bias provides insights into the cognitive abilities of tortoises.