Surfing the biomass size spectrum: some remarks on history, theory, and application

Charles Elton introduced the "pyramid of numbers" in the late 1920s, but this remarkable insight into body-size dependent patterns in natural communities lay fallow until the theory of the biomass size spectrum was introduced by aquatic ecologists in the mid-1960s. They noticed that the summed biomass concentration of individual aquatic organisms was roughly constant across equal logarithmic intervals of body size from bacteria to the largest predators. These observations formed the basis for a theory of aquatic ecosystems, based on the body size of individual organisms, that revealed new insights into constraints on the structure of biological communities. In this review, we discuss the history of the biomass spectrum and the development of underlying theories. We indicate how to construct biomass spectra from sample data, explain the mathematical relations among them, show empirical examples of their various forms, and give details on how to statistically fit the most robust linear and nonlinear models to biomass spectra. We finish by giving examples of biomass spectrum applications to production and fisheries ecology and offering recommendations to help standardize use of the biomass spectrum in aquatic ecology.