Incorporating temporally dynamic baselines in isotopic mixing models

Stable isotopes (particularly C and N) are widely used to make inferences regarding food web structure and the phenology of consumer diet shifts, applications that require accurate isotopic characterization of trophic resources to avoid biased inferences of feeding relationships. For example, most isotope mixing models require that endmembers be adequately represented by a single probability distribution; yet, there is mounting evidence that the isotopic composition of aquatic organisms often used as mixing model endmembers can change over periods of weeks to months. A review of the literature indicated that the delta13C values of five aquatic primary consumer taxa, commonly used as proxies of carbon production sources (i.e., trophic baselines), express seasonally dynamic cycles characterized by an oscillation between summer maxima and winter minima. Based on these results, we built a dynamic baseline mixing model that allows a growing consumer to track temporal gradients in the isotopic baselines of a food web. Simulations showed that the ability of a consumer to maintain or approach isotopic equilibrium with its diet over a realistic growth season was strongly affected by both the rate of change of the isotopic baseline and equilibration rate of the consumer. In an empirical application, mixing models of varying complexity were used to estimate the relative contribution of benthic vs. pelagic carbon sources to nine species of juvenile fish in a fluvial lake of the St. Lawrence River system (Québec, Canada). Estimates of p (proportion of carbon derived from benthic sources) derived from a static mixing model indicated broad interspecific variation in trophic niche, ranging from complete benthivory to > 95% reliance on pelagic food webs. Output from the more realistic dynamic baseline mixing model increased estimated benthivory by an average of 36% among species. Taken together, our results demonstrate that failing to identify dynamic baselines when present, and (or) matching consumers with baseline taxa that possess substantially different equilibration rates can seriously bias interpretation of stable isotope data. Additionally, by providing a formalized framework that allows both resources and consumers to shift their isotopic value through time, our model demonstrates a feasible approach for incorporating temporally dynamic isotope conditions in trophic studies of higher consumers.