Biology of the planktonic stages of benthic octopuses

Octopuses of the family Octopodidae adopt two major life-history strategies. The first is the production of relatively few, large eggs resulting in well-developed hatchlings that resemble the adults and rapidly adopt the benthic habit of their parents. The second strategy is production of numerous small eggs that hatch into planktonic, free-swimming hatchlings with few suckers, simple chromatophores and transparent musculature. These distinctive planktonic stages are termed para-larvae and differ from conspecific adults in their morphology, physiology, ecology and behaviour. This study aims to review available knowledge on this subject. In benthic octopuses with plank. tonic stages, spawning characteristics and duration of planktonic life seem to play an important role in their dispersal capacities. Duration of the hatching period of a single egg mass can range from 2 days to 11 wk, while duration of the planktonic stage can range from 3 wk to half a year, depending on the species and temperature. Thus these paralarvae possess considerable potential for dispersal. In some species, individuals reach relatively large sizes while living as part of the micronekton of oceanic, epipelagic waters. Such forms appear to delay settlement for an unknown period that is suspected to be longer than for paralarvae in more coastal, neritic waters. During the planktonic period, paralarval octopuses feed on crustaceans as their primary prey. In addition to the protein, critical to the protein-based metabolism of octopuses (and all cephalopods), the lipid and copper contents of the prey also appear important in maintaining normal growth. Littoral and oceanic fishes are their main predators and defence behaviours may involve fast swimming speeds, use of ink decoys, dive responses and camouflage. Sensory systems of planktonic stages include photo-, mechano- and chemoreceptors controlled by a highly evolved nervous system that follows the general pattern described for adult cephalopods. On settlement, a major metamorphosis occurs in morphology, physiology and behaviour. Morphological changes associated with the settlement process include positive allometric arm growth; chromatophore, iridophore and leucophore genesis; development of skin sculptural components and a horizontal pupillary response. At the same time, animals lose the Kolliker organs that cover the body surface, the 'lateral line system' and the oral denticles of the beaks. Strong positive phototaxis is a common response for hatchlings and some later paralarval stages but this response reduces, disappears or reverses after settlement. There are many gaps in our knowledge of the planktonic phases of benthic octopuses. Most of our understanding of octopus paralarvae comes from studies of just two species (Octopus vulgaris and Enteroctopus dofleini) and knowledge of the vast majority of benthic octopus species with planktonic stages is considered rudimentary or non-existent. Research is needed in a variety of fields, from taxonomy to ecology. Studies of feeding and nutrition are critical in order to develop the nascent aquaculture of key species and ageing studies are necessary to understand planktonic population dynamics, particularly in commercially valuable species targeted by fisheries. Current and potential anthropogenic impacts on these early life stages of octopuses, such as pollution, overfishing and global warming, are also identified.