Bet-hedging dispersal strategy of a specialist marine herbivore: a settlement dimorphism among sibling larvae of Alderia modesta

Dispersal polymorphisms are known from many terrestrial organisms that inhabit fluctuating environments, but they are not well-recognized among marine invertebrates. An unusual form of variation was found in the settlement behavior of lecithotrophic larvae of the mollusc Alderia modesta (Lovén, 1844) that markedly altered the dispersal potential of sibling larvae. Most clutches contained, in variable proportions, larvae that spontaneously metamorphosed in the egg mass or within 2 d of hatching, and larvae that delayed metamorphosis until encountering the obligate adult food, the yellow-green alga Vaucheria longicaulis. The proportion of larvae undergoing spontaneous metamorphosis within a clutch was highly variable, ranging from 0 to 90%, and was unrelated to adult size. The proportion of intracapsular metamorphosis was also variable, ranging from 0 to 32%. The percentage of spontaneous metamorphosis per clutch was phenotypically plastic: when adults were starved, there was a significant tendency for each successive clutch to have a lower percentage of spontaneous metamorphosis, thus increasing the dispersal potential of the offspring. The decrease in spontaneous metamorphosis was significant after only 24 h of starvation. There was no significant increase in the cumulative level of spontaneous metamorphosis after the 2nd day post-hatching, but larval mortality increased as a linear function after the 5th day. Most larvae that did not spontaneously settle in the first 2 d delayed metamorphosis until they were exposed to the adult host alga, V. longicaulis. When 2 d old larvae were exposed to 17 species of macroalgae and sediment from the adult habitat, only V. longicaulis induced significant metamorphosis (93.3 ± 6.7%); all other species of algae and field sediment induced no metamorphosis or low levels (0–9%) that were not significantly higher than sea water controls. There was no difference in the settlement specificity of 2 and 9 d old larvae when tested against 2 different algae, nor in their ability to complete metamorphosis. The high initial percentage of spontaneous metamorphosis reduces the dispersal potential of some larvae in each clutch relative to their siblings that delay metamorphosis until stimulated by V. longicaulis; this settlement dimorphism represents a novel bet-hedging strategy among marine invertebrates.