RESUMEN
A critical question in marine ecology is understanding how organisms will cope with environmental conditions under climate change. Increasing temperatures not only have a direct effect on marine organisms but may also lead to food limitation through for example trophic mismatches, or by the increased metabolic demands imposed by developing at high temperatures. Using barnacles from a population of North Wales, we studied the combined effect of temperature and food density on the survival, settlement success, developmental time and body size of larvae of the native barnacle Semibalanus balanoides and its exotic competitor, the barnacle Austrominius modestus. Larvae were reared at similar food levels but at temperature ranges which varied among species reflecting their different phenology and tolerances. For S. balanoides (spring larval release) we used a lower temperature of 9 °C, reflecting spring temperatures from N Wales to SW England, and 15 °C representing warmer conditions; for A. modestus (summer larval release) a typical summer temperature for this geographic range of 15 °C was used with a raised temperature of 18 °C. Larvae were reared under controlled conditions in automated, computer programmable incubators and fed diatoms (Skeletonema costatum) at three food levels. We found stress effects of food limitation on larval performance of S. balanoides. While survival during naupliar development was little affected by food and temperature, low food levels strongly depressed survival and settlement during the cyprid stage of S. balanoides at both tested temperatures, but especially at 15 °C. By contrast, at the tested temperatures little effects were found on survival and settlement success in the exotic A. modestus. Both species delayed development in response to low food levels while S. balanoides cyprids showed decreased body size at the high tested temperature. The main impact occurred as a delayed effect, at the time when cyprids attempt to settle, rather than as an effect on naupliar survival or metamorphosis to the cyprid stage. Response in body size and developmental time may have costs at the time of metamorphosis (delayed settlement) or after metamorphosis. Overall, our experiments suggest that as temperature increases, settlement success of S. balanoides larvae (but not that of its competitor A. modestus) will become more sensitive to conditions of food limitation, imposed for instance by phenological mismatches with periods of phytoplankton peak.