Adaptive diversity in congeneric coastal crabs: Ontogenetic patterns of osmoregulation match life-history strategies in Armases spp (Decapoda, Sesarmidae)

Ontogeny of osmoregulation and salinity tolerance were investigated throughout the larval development of two congeneric species of sesarmid crab, Armases ricordi (H. Milne Edwards) and A. roberti (H. Milne Edwards), and compared with previous observations from two further congeners, A. miersii (Rathbun) and A. angustipes (Dana). In the semiterrestrial coastal species A. ricordi, the zoeal stages were only at moderately reduced salinities (17-25.5‰) capable of hyper-osmoregulation, being osmoconformers at higher concentrations. The megalopa was the first ontogenetic stage of this species, which exhibited significant hyper-osmoregulation at further reduced salinities (≥ 5‰), as well as a moderately developed function of hypo-regulation at high concentrations (32-44‰). The riverine species A. roberti showed similar overall patterns in the ontogeny of osmoregulation, however, also some striking differences. In particular, its first zoeal stage showed already at hatching a strong capability of hyper-osmoregulation in salinities down to 5‰. Interestingly, this early expressed function became significantly weaker in the subsequent zoeal stages, where survival and capabilities of hyper-osmoregulation were observed only at salinities down to 10‰. The function of hyper-regulation in strongly dilute media re-appeared later, in the megalopa stage, which tolerated even an exposure to freshwater (0.2‰). Differential species- and stage-specific patterns of osmoregulation were compared with contrasting life styles, reproductive behaviours, and life-history strategies. In A. ricordi, the larvae are released into coastal marine waters, where salinities are high, and thus, no strong hyper-osmoregulation is needed throughout the zoeal phase. The megalopa stage of this species, by contrast, may invade brackish mangrove habitats, where osmoregulatory capabilities are required. Strong hyper-osmoregulation occurring in both the initial and final larval stages (but not in the intermediate zoeal stages) of A. roberti correspond to patterns of ontogenetic migration in this species, including hatching in freshwater, larval downstream transport, later zoeal development in estuarine waters, and final re-immigration of megalopae and juvenile crabs into limnic habitats, where the conspecific adults live. Similar developmental changes in the ecology and physiology of early life-history stages seem to occur also in A. angustipes. A. miersii differs from all other species, showing an early expression and a gradual subsequent increase of the function of hyper-osmoregulation. This ontogenetic pattern corresponds with an unusual reproductive biology of this species, which breeds in supratidal (i.e. land-locked) rock pools, where variations in salinity are high and unpredictable. Matching patterns in the ontogeny of osmoregulation and life-history strategies indicate a crucial adaptive role of osmoregulation for invasions of (by origin marine) crabs into brackish, limnic and terrestrial environments.