How modularity and heterotrophy complicate the understanding of the causes of thermal performance curves: the case of feeding rate in a filter-feeding animal.

Warming global temperatures have consequences for biological rates. Feeding rates reflect the intake of energy that fuels survival, growth and reproduction. However, temperature can also affect food abundance and quality, as well as feeding behavior, which all affect feeding rate, making it challenging to understand the pathways by which temperature affects the intake of energy. Therefore, we experimentally assessed how clearance rate varied across a thermal gradient in a filter-feeding colonial marine invertebrate (the bryozoan Bugula neritina). We also assessed how temperature affects phytoplankton as a food source, and zooid states within a colony that affect energy budgets and feeding behavior. Clearance rate increased linearly from 18°C to 32°C, a temperature range that the population experiences most of the year. However, temperature increased algal cell size, and decreased the proportion of feeding zooids, suggesting indirect effects of temperature on clearance rates. Temperature increased polypide regression, possibly as a stress response because satiation occurred quicker, or because phytoplankton quality declined. Temperature had a greater effect on clearance rate per feeding zooid than it did per total zooids. Together, these results suggest that the effect of temperature on clearance rate at the colony level is not just the outcome of individual zooids feeding more in direct response to temperature but also emerges from temperature increasing polypide regression and the remaining zooids increasing their feeding rates in response. Our study highlights some of the challenges for understanding why temperature affects feeding rates, especially for understudied, yet ecologically important, marine colonial organisms.