Organismal form constrains the evolution of complex lever systems in Neotropical cichlid four-bar linkages.

The diversification of functional traits may be limited by the intrinsic constraints of organismal form (i.e., constructional constraints), owing to the differential investment in different anatomical structures. In this study, we test whether overall organismal form impacts the evolution of shape and function in complex lever systems. We examined the relationship between four-bar shape and overall head shape in two four-bar linkage systems: the oral-jaw and hyoid-neurocranium systems in Neotropical cichlids. We also investigated the strength of form-function mapping in these four-bar linkages and the impact of constraining head shape on these correlations. We quantified the shape of the head and two four-bar linkages using geometric morphometrics and compared these with the kinematic transmission coefficient of each linkage system. The shapes of both linkages were strongly correlated with their mechanical properties, and head shape appears to constrain the shape of both four-bar linkages. Head shape induced greater integration between the two linkages, was associated with stronger form-function correlations and higher rates of evolution in biomechanically important features. Head shape constraints may also contribute to a weak but significant trade-off in linkage kinematics. Elongation of the head and body, in particular, appears to minimize the impact of this trade-off, possibly through maximizing anterior-posterior space availability. However, the strength of relationships between shape and function, and the impact of head shape differed between the two linkages, with the hyoid four-bar in general showing stronger form-function relationships despite being more independent from head shape constraints.