The settlement stage in the common octopus Octopus vulgaris Cuvier, 1797: a complex transition between planktonic and benthic lifestyles

Merobenthic octopods like Octopus vulgaris undergo a transitional period between the planktonic and benthic phases, known as settlement stage. In this work, three sub-stages (“pre-settlement”, “settlement” and “post-settlement”) have been defined based on morphological, anatomical, and behavioural changes. At the end of the planktonic phase advanced paralarvae are transparent with 65–80 chromatophores, iridophores covering eyes and digestive system, Kölliker organs, circular pupils, ~ 20 suckers per arm (spa), and mantle length (ML) bigger than total length (TL; ML/TL > 60%). The “pre-settlement” sub-stage (ML/TL from 65 to 55%, ~ 20–25 spa) is marked by the onset of clinging reflexes, where the pre-settlers touch the walls and bottom of the tank and start crawling clumsily. Morphologically, they are transparent with increased chromatophores along the arms and iridophores around the eyes and head. During the “settlement” sub-stage (~ 55–48% ML/TL, ~ 25–35 spa) there is an exponential increase of chromatophores in the dorsal area of head and mantle, and the settlers show strong negative phototaxis, crawling for shelter when disturbed. The skin is still transparent but new chromatic cells (leucophores) develop and Kölliker organs are almost lost. During the “post-settlement” sub-stage (~ 48–40% ML/TL, > 35 spa) the post-settlers display very fast and coordinated movements, have horizontal pupils, and develop the “eye-bar”. The chromatic cells keep increasing exponentially, giving a pale appearance to the skin. The beginning of the benthic phase in O. vulgaris juveniles is marked by the presence of skin sculptural components (papillae) and a complex display of chromatic, postural, and cryptic patterns.