Water temperature and season length interact to explain a rare non-linear ecogeographic cline in body size

Aim: Adult body size often exhibits patterns across large-scale environmental gradients, creating ecogeographic clines. However, the form of body size clines varies across taxonomic groups, with linear and non-linear patterns in body size observed in nature. Non-linear body size clines have received less study, and questions remain about how environmental gradients interact to produce non-linear clines. We examined the body size of the American horseshoe crab (Limulus polyphemus), a widely distributed marine arthropod, and evaluated the hypothesis that temperature and active season length can interact multiplicatively to result in a dome-shaped distribution.Location: Fourteen states in the United States of America and three Mexican states, representing the entire geographic range of the species.Methods: We compiled environmental data and body size measurements from more than 49,000 individual horseshoe crabs. For each location, we extracted from the literature or calculated from raw data the mean male prosoma width and the mean female prosoma width. We applied a general additive modelling (GAM) approach to characterize the body size cline, test a hypothesis regarding temperature and season length, and explore evidence for the influence of additional environmental factors.Results: Model results indicate temperature and season length could act multiplicatively to produce dome-shaped clines, and these findings align with and quantify previous anecdotal reports of a strong dome-shaped body size cline across latitude for horseshoe crabs.Main Conclusions: Active season length appears to become relatively more influential on horseshoe crab body size in the northern part of their range, while temperature effects per se appear to dominate in southern latitudes. For horseshoe crabs, the pattern of size variation is consistent with the predictions of Optimal Resource Allocation models, but more study is needed to elucidate mechanistic underpinnings. Considering climate change projections, results from our study suggest future shifts in horseshoe crab body sizes.