Complex patterns of expression of heat-shock protein 70 across the southern biogeographical ranges of the intertidal mussel Mytilus californianus and snail Nucella ostrina

Aim It is often assumed that species reach their highest densities in the centre of their ranges and decline in abundance toward the edges of the range. Implicit in this notion, which we call the abundant centre hypothesis, is the assumption that the edges of the range are more stressful to organisms and are more likely to show responses to climate change. However, an earlier review and empirical study of patterns of abundance across the range of intertidal invertebrates show little support for the abundant centre hypothesis and further demonstrated that few studies have examined patterns in either abundance or stress across species ranges. In part this gap is due to the logistical difficulties of sampling species across large geographical ranges. Here we use intertidal invertebrates, which have relatively simple linear latitudinal ranges, and heat-shock proteins, which have been shown to be an integrative measure of organismal stress, to test the hypothesis that species are more stressed at the edges of their range. We use complementary data on population density to test the relationship between stress proteins and overall species density across the species’ range. Location Our sampling programme covered the southern half of the large geographical ranges of two intertidal invertebrates on the Pacific Coast of North America. Sites were spread between northern Baja California, Mexico and Vancouver Island, Canada, a range of c. 22 degrees of latitude. Method We sampled levels of heat-shock protein 70 (Hsp70) in eight to 12 individuals from each of 20 sites for the intertidal mussel Mytilus californianus and 11 sites for the intertidal snail Nucella ostrina, spread throughout the southern half of their geographical ranges. The relationships between levels of Hsp70 in individuals from a site and (1) latitude of the site, (2) the site's position in the species’ range and (3) average population density were determined. Results No significant relationship was found in either species between levels of Hsp70 and latitude, position in the range or population density. Complex patterns that did emerge may be explained by nonlinear gradients in environmental conditions along the Pacific coast. Specifically, we observed peak values of Hsp70 for both species in northern Oregon, where intertidal zones are disproportionately exposed to daytime emersion (exposure to air) in the summer months of collections. A second peak for M. californianus was found south of Point Conception, California, which is marked by dramatic shifts toward warmer sea temperatures and decreased wave exposure. Main conclusions Patterns that emerged were not predicted by simple models based on the abundant centre hypothesis. However, they are consistent with more complex pictures of heat stress, organismal condition and abundance along a latitudinal gradient that have been demonstrated in recent studies. We suggest that latitudinal complexity, species-specific differences and local effects must be considered before generalizing the relationship between environmental stress, abundance, range limits and responses of ranges to climate change.