Morphological comparison of the cranial movement apparatus in mudskippers (Gobiidae: Oxudercinae)

Possession of the neck allows vertebrates to move the head independently from the trunk. Fish do not have the neck and thus the cranial mobility could be limited. Oxudercine gobies show full range of habitat transition from aquatic to terrestrial environments and exhibit flexible cranial movement, yet the cranium-movement apparatus is little known. In this study, we investigated the anatomy of the structure of the eight oxudercine gobies, Oxuderces nexipinnis, Parapocryptes serperaster, Pseudapocryptes elongatus, Scartelaos histophorus, Boleophthalmus boddarti, Periophthalmus chrysospilos, Periophthalmodon schlosseri, and Periophthalmodon septemradiatus. These species share similarities in the specialized features of the craniovertebral joint and the epaxials attaching onto different locations of the neurocranium. On the other hand, large space between the ventral portions of the craniovertebral joint only occurs in O. nexipinnis, Pd. elongatus, Pn. schlosseri and Pn. septemradiatus. Hypaxials are hypertrophied at the insertion point and attach more anteriorly onto the ventral side of the neurocranium in B. boddarti, O. nexipinnis, Pa. serperaster, Pd. elongatus, and S. histophorus, whereas the muscles are small and attach posteriorly in the remaining species. There were significant differences in the area occupancy ratio of the post-cranial neural spines, the lever arm ratio of the cranial rotation, and the angle between the horizontal plane and the plane through the craniovertebral joint among the species. The cranial depression presumably facilitates grazing of oxudercine gobies in their early stage of terrestrial transition, whereas the cranial elevation parameters are contradictory to the terrestrial gradient. The cranium-movement morphometrics partially agree with the phylogeny.