Cranial musculature of batoids: A standardized nomenclature

Batoids (rays and skates) are cartilaginous fishes whose jaws are not articulated directly to the neurocranium. The only point of contact between them are the hyomandibular cartilages, resulting in a unique mandibular suspension called euhyostyly. Due to this decoupling of the jaws from the skull, muscles play an essential role in modulating mandibular movements during the feeding process, especially during mandibular protrusion. The main objectives of our study were: (1) to examine the mandibular musculature of eight batoid species from different orders in the Batoidea and (2) establish a standardized musclulature terminology for future comparative myological studies in batoids. For each muscle bundle, the general characteristics of each cranial muscle were described and their origin and insertions were identified. The number of muscle bundles differed intraspecifically. On the dorsal surface, we reported the first evidence of the presence of the precranial muscle (PCM) in U. halleri, as well as the ethmoideo-parethmoidalis muscle (ETM) in R. velezi, P. glaugostigma and Z. exasperata; in addition, the insertion of the spiracularis muscle (SP) extended to the ventral surface of the oropharyngeal tract in myliobatiforms. On the ventral surface of the head, both N. entemedor and M. californica exhibited additional muscles in the mandibular area. These muscles were renamed as part of the standardization of mandibular terminology: the depressor mandibularis minor (DMM) in N. entemedor and the adductor mandibulae profundus (AMP) in M. californica. The standardization of terminology is essential for futures studies of the mandibular apparatus in batoids, to facilitate the morphological description of muscles in species without anatomical accounts and for continuity in broader comparative analyses.