Delayed metamorphosis in decapod crustaceans: evidence and consequences Retraso de la metamorfosis en crustáceos decápodos: evidencias y consecuencias

Most marine invertebrate species exhibit a complex life cycle including a planktonic larval phase and a benthic juvenileadult phase. Metamorphosis and settlement are the links between these phases of development. In many species, metamorphosis is triggered by specific chemical and/or physical cues, mainly associated with the adult habitat. In the absence of such cues, competent larvae can delay their metamorphosis by a few days to several months. Most investigations on the delay of metamorphosis have been realised on sessile or sedentary species. In relation to mobile decapod crustaceans, the number of such studies is low, probably because the members of this group retain their mobility after metamorphosis, and hence, may depend less on enviromental cues for the induction of settlement and metamorphosis. Nevertheless, the larvae of some decapod species have been shown to depend on metamorphosis-stimulating cues. These include special types of substrates, physical or chemical traits of particular (e.g., estuarine) water bodies, as well as odors from conspecific or congeneric adults. The capacity for delay is, in the decapod species studied so far, limited and may normally end with spontaneous metamorphosis. An extended time of larval development presents the advantage of enhancing the probability for locating a suitable habitat, but it may imply, as a disadvantage, a reduction of juvenile growth or survival and a prolonged development time preceding benthic life. This paper reviews the available evidence for delayed metamorphosis in decapod crustaceans, indentifed cues, the importance of larval age at the time of contact with a cue, and costs of delayed metamorphosis. Additionally, we propose new frontiers for future investigations on delayed metamorphosis in decapod crustaceans, including the molecular identification of chemical cues, the identification of the stage(s) of the moulting cycle that is or are sensitive to such cues, the study of hormonally mediated effects on the moulting cycle, the quantification of energetic or other costs of delayed metamorphosis, and the analysis of relationships between the effectiveness of adult odors and phylogenetic proximity of larvae and adults.