Chronobiology of deep-water decapod crustaceans on continental margins

Species have evolved biological rhythms in behaviour and physiology with a 24-h periodicity in order to increase their fitness, anticipating the onset of unfavourable habitat conditions. In marine organisms inhabiting deep-water continental margins (i.e. the submerged outer edges of continents), day-night activity rhythms are often referred to in three ways: vertical water column migrations (i.e. pelagic), horizontal displacements within benthic boundary layer of the continental margin, along bathymetric gradients (i.e. nektobenthic), and endobenthic movements (i.e. rhythmic emergence from the substrate). Many studies have been conducted on crustacean decapods that migrate vertically in the water column, but much less information is available for other endobenthic and nektobenthic species. Also, the types of displacement and major life habits of most marine species are still largely unknown, especially in deep-water continental margins, where steep clines in habitat factors (i.e. light intensity and its spectral quality, sediment characteristics, and hydrography) take place. This is the result of technical difficulties in performing temporally scheduled sampling and laboratory testing on living specimens. According to this scenario, there are several major issues that still need extensive research in deep-water crustacean decapods. First, the regulation of their behaviour and physiology by a biological clock is almost unknown compared to data for coastal species that are easily accessible to direct observation and sampling. Second, biological rhythms may change at different life stages (i.e. size-related variations) or at different moments of the reproductive cycle (e.g. at egg-bearing) based on different intra- and interspecific interactions. Third, there is still a major lack of knowledge on the links that exist among the observed bathymetric distributions of species and selected autoecological traits that are controlled by their biological clock, such as the diel rhythm of behaviour. Species evolved in a photically variable environment where intra- and inter-specific interactions change along with the community structure over 24 h. Accordingly, the regulation of their biology through a biological clock may be the major evolutionary constraint that is responsible for their reported bathymetric distributions. In this review, our aim is to propose a series of innovative guidelines for a discussion of the modulation of behavioural rhythms of adult decapod crustaceans, focusing on the deep waters of the continental margin areas of the Mediterranean as a paradigm for other marine zones of the world.