Swimming with multiple propulsors: measurement and comparison of swimming gaits in three species of neotropical cichlids.

Comparative studies of fish swimming have been limited by the lack of quantitative definitions of fish gaits. Traditionally, steady swimming gaits have been defined categorically by the fin or region of the body that is used as the main propulsor and named after major fish clades (e.g. carangiform, anguilliform, balistiform, labriform). This method of categorization is limited by the lack of explicit measurements, the inability to incorporate contributions of multiple propulsors and the inability to compare gaits across different categories. I propose an alternative framework for the definition and comparison of fish gaits based on the propulsive contribution of each structure (body and/or fin) being used as a propulsor relative to locomotor output, and demonstrate the effectiveness of this framework by comparing three species of neotropical cichlids with different body shapes. This approach is modular with respect to the number of propulsors considered, flexible with respect to the definition of the propulsive inputs and the locomotor output of interest, and designed explicitly to handle combinations of propulsors. Using this approach, gait can be defined as a trajectory through propulsive space, and gait transitions can be defined as discontinuities in the gait trajectory. By measuring and defining gait in this way, patterns of clustering corresponding to existing categorical definitions of gait may emerge, and gaits can be rigorously compared across categories.