The development of orofacial complex in bats: Implications for orofacial clefting

Orofacial morphology in mammals plays a critical role in essential life functions such as feeding and communication, which are influenced by the shapes of these anatomical structures. Bats are known to exhibit highly diversified orofacial morphotypes within their clade, reflecting their varied diets and echolocation behaviors. The presence of bony discontinuities between the premaxilla and maxilla or among the premaxillae is a notable feature of bat orofacial morphology, observed in certain lineages. It is suggested that these unique orofacial morphotypes, not generally found in other mammals, have evolved in relation to dietary adaptations rather than merely for echolocation mode. Until now, the developmental background of the bony discontinuities in the bat orofacial complex has been insufficiently investigated. Here, we present a comparative study of the chondrocranium and epithelial organs in the orofacial complex of three bat species: Cynopterus sphinx, Rhinolophus malayanus, and Vespertilio sinensis. Our observations indicate that the preceding morphogenesis of orofacial cartilage and epithelial structures is remarkably different among these three species. In C. sphinx and V. sinensis, the region forming from the regression of the palatine process of the premaxilla was filled with orofacial cartilage and epithelial structures. We also found that the clefted morphology observed in R. malayanus and V. sinensis was formed via contrastingly divergent developmental processes. Midline clefts among Yangochiroptera have been previously categorized to represent a uniform morphotype, but our study highlights that attributing midline clefts into a singular category should be revisited, advocating for a nuanced categorization of cleft morphology based on their morphogenetic patterns. Further research on the bat orofacial complex may enhance our understanding of bat evolutionary diversification and offer insights into the developmental mechanisms of human cleft palate.