Fish collective behaviours provide several benefits to conspecific individuals, although mixed-species aggregations have been reported to often occur. However, the mechanisms promoting phenotypically heterogeneous fish aggregations have been poorly explored so far. Herein, the neon tetra Paracheirodon innesi was selected as ideal model organism to test the role of visible phenotypic traits in promoting fish shoaling. Robotic fish replicas of different colour (e.g. biomimetic livery, blue livery, red livery, grey livery), but with the morphology inspired to 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 replica, the blue replica, the red replica and the grey replica. This could be due to the greater visibility of the blue colour even in dark conditions in these fish. Furthermore, an increased reddening of the livery is often caused by physiological processes related to a non-optimal 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 presence of food. This confirms the hypothesis that heterogeneous shoals are promoted by the anti-predator benefits, and reduced by competition. Our animal-robot interaction study allowed to understand basic features of the behavioural ecology favouring heterogeneous aggregations in shoaling fish, as well as provided a novel paradigm, based on biohybridization, for the artificial life synthetic methods.

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