Living on the upper intertidal mudflat: Different behavioral and physiological responses to high temperature between two sympatric cerithidea snails with divergent habitat-use strategies.

Temperature plays a major role in controlling species' distributions, and small-scale variation in the thermal environment are potentially an important factor that governs distributions on a local scale. For untangling the roles of behavioral and physiological adaptations on species' distribution at a small-scale level, we carried out a comparative study of two mudflat snails (genus Cerithidea) by determining these congeners' burying behavior, lethal temperature, cardiac performance and heat-shock protein (hsp70) gene expression. These two sympatric snails occupy different microhabitats on the upper intertidal mudflat. During periods of emersion, C. cingulata inhabits the open mudflat and C. largillierti usually aggregates around small rocks on the upper intertidal mudflat. Our results indicate that the two Cerithidea congeners show different behavioral and physiological responses to high temperature. Compared to C. largillierti, C. cingulata prefers to bury into the mud, has a higher thermal limit and a higher level of inducible expression of hsp70 mRNA, implying important roles of behavioral and physiological adaptations to the harsh thermal environment on the open mudflat. Furthermore, results of generalized additive modelling (GAM) analysis of cardiac performance and coefficient of variation (CV) of hsp70 mRNA expression showed high inter-individual variation in C. cingulata. These results highlight the importance of behavioral and physiological adaptions in sympatric species' distributions on the mudflat and help to shed light on the mechanisms of how small-scale differences in the thermal environment shape sympatric species' distributions.