Delimiting the boundaries of sesamoid identities under the network theory framework

Sesamoid identity has long been the focus of debate, and how they are linked to other elements of the skeleton has often been considered relevant to their definition. A driving hypothesis of our work was that sesamoids' nature relies deeply on their connections, and thus we propose an explicit network framework to investigate this subject in Leptodactylus latinasus (Anura: Leptodactylidae). Through the dissection of L. latinasus' skeleton, we modeled its anatomical network where skeletal elements were considered nodes while joints, muscles, tendons, and aponeurosis were considered links. The skeletal elements were categorized into canonical skeletal pieces, embedded sesamoids, and glide sesamoids. We inquired about the general network characterization and we have explored further into sesamoid connectivity behavior. We found that the network is structured in a modular hierarchical organization, with five modules on the first level and two modules on the second one. The modules reflect a functional, rather than a topological proximity clustering of the skeleton. The 25 sesamoid pieces are members of four of the first-level modules. Node parameters (centrality indicators) showed that: (i) sesamoids are, in general terms, peripheral elements of the skeleton, loosely connected to the canonical bone structures; (ii) embedded sesamoids are not significantly distinguishable from canonical skeletal elements; and (iii) glide sesamoids exhibit the lowest centrality values and strongly differ from both canonical skeletal elements and embedded sesamoids. The loose connectivity pattern of sesamoids, especially glides, could be related to their evolvability, which in turn seems to be reflected in their morphological variation and facultative expression. Based on the connectivity differences among skeletal categories found in our study, an open question remains: can embedded and glide sesamoids be defined under the same criteria? This study presents a new approach to the study of sesamoid identity and to the knowledge of their morphological evolution.