Any colour you like: fish interacting with bioinspired robots unravel mechanisms promoting mixed phenotype aggregations

Collective behaviours in homogeneous shoals provide several benefits to conspecifics, although mixed-species aggregations have been reported to often occur. Mixed aggregations may confer several beneficial effects such as antipredator and foraging advantages. However, the mechanisms promoting phenotypically heterogeneous fish aggregations have been poorly explored so far. Herein, the neon tetra Paracheirodon innesi was selected as the ideal model organism to test the role of visible phenotypic traits in promoting fish shoaling. Robotic fish replicas of different colours, but with a morphology inspired by P. innesi, were developed to test the affiliation behaviour of neon tetra individuals towards fish replicas with different phenotypic traits. P. innesi individuals showed a decreasing preference in shoaling with the biomimetic, the blue, the red, and the grey replicas. This could be due to the greater visibility of the blue colour even in dark conditions. Furthermore, an increased reddening of the livery is often caused by physiological processes related to a nonoptimal behavioural status. The time spent in shoaling with each fish replica was strongly influenced by different ecological contexts. The longest shoaling duration was observed when a biomimetic predator was present, while the shortest shoaling duration was recorded in the presence of food. This confirms the hypothesis that heterogeneous shoals are promoted by the antipredator benefits, and reduced by competition. This study allowed us to understand basic features of the behavioural ecology favouring heterogeneous aggregations in shoaling fish, and provided a novel paradigm for biohybrid robotics.