Nudibranchs out of water: long-term temporal variations in the abundance of two Dendrodoris species under emersion

The sudden appearance and disappearance of nudibranchs in intertidal areas have puzzled researchers all over the world, giving rise to a great diversity of theories to explain it. Here we conducted a five-year survey to evaluate seasonal changes in the abundance of Dendrodoris herytra and D. grandiflora in the Sado estuary (Portugal) and to explore a possible relationship with environmental factors such as temperature, salinity, turbidity and dissolved oxygen. Moreover, we report, for the first time, the capacity of Dendrodoris nudibranchs to tolerate emersion (unhidden and completely exposed to sun exposure) during low tides. Our results showed that both species consistently started to appear emerged in March, reaching a peak abundance between April and May, and completely disappearing in July. In both species, this temporal trend was significantly associated with water temperature, turbidity, and dissolved oxygen, but not with salinity. We argue that the sudden appearance and disappearance of these nudibranchs in intertidal areas may result from a seasonal horizontal movement of adult nudibranchs from subtidal areas to mate in intertidal areas during spring, when phytoplankton production is enhanced and planktotrophic larvae may benefit from greater food availability.