A Dynamic Energy Budget (DEB) model for the keystone predator Pisaster ochraceus

We present a Dynamic Energy Budget (DEB) model for the quintessential keystone predator, the rocky-intertidal sea star Pisaster ochraceus. Based on first principles, DEB theory is used to illuminate underlying physiological processes (maintenance, growth, development, and reproduction), thus providing a framework to predict individual-level responses to environmental change. We parameterized the model for P. ochraceus using both data from the literature and experiments conducted specifically for the DEB framework. We devoted special attention to the model's capacity to (1) describe growth trajectories at different life-stages, including pelagic larval and post-metamorphic phases, (2) simulate shrinkage when prey availability is insufficient to meet maintenance requirements, and (3) deal with the combined effects of changing body temperature and food supply. We further validated the model using an independent growth data set. Using standard statistics to compare model outputs with real data (e.g. Mean Absolute Percent Error, MAPE) we demonstrated that the model is capable of tracking P. ochraceus' growth in length at different life-stages (larvae: MAPE = 12.27%; post-metamorphic, MAPE = 9.22%), as well as quantifying reproductive output index. However, the model's skill dropped when trying to predict changes in body mass (MAPE = 24.59%), potentially because of the challenge of precisely anticipating spawning events. Interestingly, the model revealed that P. ochraceus reserves contribute little to total biomass, suggesting that animals draw energy from structure when food is limited. The latter appears to drive indeterminate growth dynamics in P. ochraceus. Individual-based mechanistic models, which can illuminate underlying physiological responses, offer a viable framework for forecasting population dynamics in the keystone predator Pisaster ochraceus. The DEB model herein represents a critical step in that direction, especially in a period of increased anthropogenic pressure on natural systems and an observed recent decline in populations of this keystone species.