The urohyal: Development and homology within osteichthyans

The formation of the unpaired structure ventral to the basibranchial region, the so-called urohyal, differs within osteichthyans. A cartilaginous preformed, unpaired "urohyal" is present in sarcopterygians. A three-tendon ossification is present in Polypterus. An "urohyal" or urohyal is absent in both Amia and Lepisosteus. The urohyal formed as an unpaired ossification of the tendon of the sternohyoideus muscle is a feature of teleosts. A new structure, the parurohyal, arises as a double ossification of the tendon of the sternohyoideus muscle in siluroids; during ontogeny an anterodorsal crest or cup-like structure derives from the anterior basibranchial region and the tendon bone; therefore, the parurohyal is compound in origin. Judging from their formation and their distribution within osteichthyans the cartilaginous preformed "urohyal" and the teleostean urohyal are nonhomologous, whereas the urohyal and parurohyal are homologous. The urohyal is connected by ligaments with the ventral hypohyals in most teleosts, whereas it articulates with the ventral hypohyals in teleosts such as Anguilla and Chanos. The parurohyal is a synapomorphy of siluroids. The parurohyal in siluroids is articulated with both ventral and dorsal hypohyals, and with the basibranchial region in catfishes such as diplomystids and ictalurids, whereas it articulates only with the ventral hypohyals in other catfishes such as trichomycterines. The passage of the hypobranchial artery through the hypobranchial foramen of the parurohyal is a unique feature of siluroids, like the absence of the basihyal bone. An ossified dorsal hypohyal appears late in ontogeny in Amia, as do tooth plates related to the medial side of the hyoid arch and branchiostegal rays in Amia, and tooth plates on the hyoid arch and branchiostegal rays in Elops (unique features within extant teleosts). Two ossified hypohyals, a synapomorphy of teleosts, are present early in ontogeny. There is intraspecific variation in the onset of ossification of the bones of the head, but the sequence of ossification between bones in a defined structural system is conserved (e.g., branchiostegal rays ossify first, then bones of the hyoid arch).