Cleaner wrasse pass the mark test. What are the implications for consciousness and self-awareness testing in animals?

The ability to perceive and recognise a reflected mirror image as self (mirror self-recognition, MSR) is considered a hallmark of cognition across species. Although MSR has been reported in mammals and birds, it is not known to occur in any other major taxon. A factor potentially limiting the ability to test for MSR is that the established assay for MSR, the mark test, shows an interpretation bias towards animals with the dexterity (or limbs) required to touch a mark. Here, we show that the cleaner wrasse fish, Labroides dimidiatus, passes through all phases of the mark test: (i) social reactions towards the reflection, (ii) repeated idiosyncratic behaviours towards the mirror (contingency testing), and (iii) frequent observation of their reflection. When subsequently provided with a coloured tag, individuals attempt to remove the mark in the presence of a mirror but show no response towards transparent marks, or to coloured marks in the absence of a mirror. This remarkable finding presents a challenge to our interpretation of the mark test - do we accept that these behavioural responses in the mark test, which are taken as evidence of self-recognition in other species, mean that fish are self-aware? Or do we conclude that these behavioural patterns have a basis in a cognitive process other than self-recognition? If the former, what does this mean for our understanding of animal intelligence? If the latter, what does this mean for our application and interpretation of the mark test as a metric for animal cognitive abilities?