The evolution of pharyngeal bones and teeth in Gobioninae fishes (Teleostei: Cyprinidae) analyzed with phylogenetic comparative methods

The subfamily Gobioninae is a subgroup in the specious fish family Cyprinidae, which bears high diversity in morphological and ecological dimensions and has its most components distributed in East Asia. In this study, the pharyngeal bones and teeth of 39 species belonging to 19 genera of the Gobioninae were examined, with the phylogenetic comparative method (PCM) and correlation methods employed to analyze the character evolution. Three characters on pharyngeal bones (shape of the pharyngeal bones, extension for attachment of the pharyngo-cleithralis internus posterior (PCIP) muscle, and teeth-bearing area) and six characters of pharyngeal teeth (shape of the five teeth in the main row, number of rows of the teeth) were identified and compared. When the character states were mapped on a molecular phylogenetic tree, it was found that, to adapt to different masticatory operations, different Gobioninae species have various morphological types of pharyngeal bones and teeth: some have intermediate pharyngeal bones bearing multiple rows of diverse teeth (conical, coarsely compressed, and compressed), others have broad pharyngeal bones bearing a single row of molar teeth, and still others have narrow pharyngeal bones bearing a single row of extremely compressed teeth. Tests on the phylogenetic signal and evolutionary associations revealed that evolution of the examined characters was all phylogenetically constrained and correlated. Owing to the homoplasy in evolution, it was suggested that the conventional method of using pharyngeal bones and teeth for phylogenetic reconstruction of cyprinid fishes should not be encouraged.