A vertebral column consisting of a persistent notochord and ossified arcocentra is the primitive condition for Gnathostomata; it still persists in primitive actinopterygians and sarcopterygians. Advanced actinopterygians and sarcopterygians develop numerous types of centra that include, among others, the presence of holocentrum, chordacentrum, and autocentrum. The chordacentrum, a mineralization or calcification of the fibrous sheath of the notochord, is only found in actinopterygians, whereas an autocentrum is a synapomorphy of teleosts above Leptolepis coryphaenoides. The chordacentrum, formed by migration of cartilaginous cells from the arches into the fibrous sheath of the notochord and usually covered by a thin calcification, is a unique feature of chondrichthyans. The actinopterygian chordacentrum and the chondrichthyan chordacentrum are not homologous. The postcaudal cartilaginous centrum is only known in postcaudal vertebrae of living dipnoans. The holocentrum is present in certain fossil dipnoans and actinopterygians, where it has been independently acquired. It is formed by proliferation of cartilage cells around the elastica externa of the notochord. These cells later ossify, forming a compact centrum. A vertebral column formed by a persistent notochord without vertebral centra is the primitive pattern for all vertebrates. The formation of centra, which is not homologous among vertebrate groups, is acquired independently in some lineages of placoderms, most advanced actinopterygians, and some dipnoans and rhipidistians. Several series of structures are associated with the vertebral column such as the supraneurals, interhaemals, radials, and ribs. In living dipnoans median neural spine, "supraneural," and dorsal radial result from growth and distal differentiation of one median cartilage into two or three median bones during ontogeny. The median neural spine articulates with the neural arch and fuses with it in the caudal vertebrae early in ontogeny. Two bones differentiate in the anterior abdominal vertebrae, i.e., the proximal neural spine and the distal "supraneural." Three bones differentiate in front of the dorsal fin, i.e., the proximal neural spine, the middle "supraneural", and the distal radial; the same pattern is observed in front of the anal fin (the proximal haemal spine, the middle interhaemal, and the distal radial). Considering that the three dorsal (and also the three ventral) bones originate from growth of only one cartilage, they cannot be serial homologs of the neural spines, or "supraneural." They are linear homologs of the median neural cartilage in living dipnoans. The development of these elements differs within osteichthyans from sarcopterygians to actinopterygians, in which the neural spine originates as a continuation of the basidorsal arcualia and in which the supraneural and radial originate from independent cartilages that appear at different times during early ontogeny. The ribs of living dipnoans are unique in that they are not articulated with parapophyses, like in primi...

The ontogenetic development of caudal vertebrae and associated skeletal elements of salmonids provides information about sequence of ossification and origin of bones that can be considered as a model for other teleosts. The ossification of elements forming the caudal skeleton follows the same sequence, independent of size and age at first appearance. Dermal bones like principal caudal rays ossify earlier than chondral bones; among dermal bones, the middle principal caudal rays ossify before the ventral and dorsal ones. Among chondral bones, the ventral hypural 1 and parhypural ossify first, followed by hypural 2 and by the ventral spine of preural centrum 2. The ossification of the dorsal chondral elements starts later than that of ventral ones. Three elements participate in the formation of a caudal vertebra: paired basidorsal and basiventral arcocentra, chordacentrum, and autocentrum; appearance of cartilaginous arcocentra precedes that of the mineralized basiventral chordacentrum, and that of the perichordal ossification of the autocentrum. Each ural centrum is mainly formed by arcocentral and chordacentrum. The autocentrum is irregularly present or absent. Some ural centra are formed only by a chordacentrum. This pattern of vertebral formation characterizes basal teleosts and primitive extant teleosts such as elopomorphs, osteoglossomorphs, and salmonids. The diural caudal skeleton is redefined as having two independent ural chordacentra plus their arcocentra, or two ural chordacentra plus their autocentra and arococentra, or only two ural chordacentra. A polyural caudal skeleton is identified by more than two ural centra, variably formed as given for the diural condition. The two ural centra of primitive teleosts may result from early fusion of ural centra 1 and 2 and of ural centra 3 and 4, or 3, 4, and 5 (e.g., elopomorphs), respectively. The two centra may corespond to ural centrum 2 and 4 only (e.g., salmonids). Additionally, ural centra 1 and 3 may be lost during the evolution of teleosts. Additional ural centra form late in ontogeny in advanced salmonids, resulting in a secondary polyural caudal skeleton. The hypural, which is a haemal spine of a ural centrum, results by growth and ossification of a single basiventral ural arococentrum and its haemal spine. The proximal part of the hypural always includes part of the ventral ural arcocentrum. The uroneural is a modification of a ural neural arch, which is demonstrated by a cartilaginous precursor. The stegural of salmonids and esocids originates from only one paired cartilaginous dorsal arcocentrum that grows anteriorly by a perichondral basal ossification and an anterodorsal membranous ossification. The true epurals of teleosts are detached neural spines of preural and ural neural arches as shown by developmental series; they are homologous to the neural spines of anterior vertebrae. Free epurals without any indication of connection with the dorsal arococentra are considered herein as an advanced state of the epural. Caudal distal radials originate fr...

The palatoquadrate and associated dermal bones have significant evolutionary transformations among teleostomes and provide numerous features that characterize teleostomian subgroups. The palatoquadrate forms the upper part of the mandibular arch and is present as a single cartilaginous element in the early ontogeny of teleostomes, except for some advanced teleosts such as siluroids where it is divided into pars autopalatina and pars pterygoquadrata. During ontogeny, the palatoquadrate may ossify as a unit, with a pars autopalatina (absent in Acanthodii), pars quadrata, and pars metapterygoidea in teleostomes (e.g., primitive acanthodians and actinopterygians, onychodonts, and rhipidistians). However, the palatoquadrate may remain cartilaginous (e.g., chondrosteans) or it may ossify as separate elements (e.g., autopalatine, metapterygoid, and quadrate) as occurs in advanced acanthodians, Polypterus and advanced actinopterygians, and advanced actinistians. From the single-unit pattern, separate autopalatine, metapterygoid, and quadrate evolve in parallel in the three teleostomian subgroups. Therefore, it is necessary to distinguish between actinopterygian and actinistian autopalatines and among acanthodian, actinopterygian, and actinistian metapterygoids and quadrates. A palatoquadrate fused with the neurocranium occurs in parallel in dipnoans. There are differences in the timing of ossification of the autopalatine, metapterygoid, and quadrate. The autopalatine ossifies late in ontogeny in Polypterus, Amia, and primitive teleosts (absent in lepisosteids and osteoglossmorphs), whereas both metapterygoid and quadrate ossify early in ontogeny. The early ossification of the autopalatine is characteristic of clupeocephalan teleosts. During ontogeny, tooth plates (not forming a separate dermometapterygoid) fuse with the metapterygoid in actinopterygians. Pars autopalatina, pars metapterygoidea, and pars quadrata are regions at the three corners of the single-unit palatoquadrate present in primitive teleostomes; there are no clear limits among these regions, but they may be identified by their processes, articular facets, and topographical relationships with surrounding bones and the orbit. Autopalatine, metapterygoid, and quadrate are chondral bones, perichondrally ossified. Dermal elements such as dermopalatine(s), entopterygoid, ectopterygoid, and tooth plates may cover the palatoquadrate medially. The predermopalatine that originates in front of pars autopalatina in Cladistia and the "dermopalatine" that lies medial to the ectopterygoid in Ginglymodi are specializations of these groups. A dermopalatine fused with the autopalatine is characteristic of clupeocephalan teleosts. Highly specialized tendon bone pterygoids are found in some teleosts (e.g., siluroids). The presence of both maxilla and lacrimal lateral to the pars autopalatina is synapomorphous of osteichthyans. The eye supported by the bony palatoquadrate is a teleostomian synapomorphy. Dermal elements support the eye in actinopterygians, the entopterygoid in...