Finding the neck-trunk boundary in snakes: anteroposterior dissociation of myological characteristics in snakes and its implications for their neck and trunk body regionalization.

The neck and trunk regionalization of the presacral musculoskeletal system in snakes and other limb-reduced squamates was assessed based on observations on craniovertebral and body wall muscles. It was confirmed that myological features characterizing the neck in quadrupedal squamates (i.e., squamates with well-developed limbs) are retained in all examined snakes, contradicting the complete lack of the neck in snakes hypothesized in previous studies. However, the posterior-most origins of the craniovertebral muscles and the anterior-most bony attachments of the body wall muscles that are located at around the neck-trunk boundary in quadrupedal squamates were found to be dissociated anteroposteriorly in snakes. Together with results of a recent study that the anterior expression boundaries of Hox genes coinciding with the neck-trunk boundary in quadrupedal amniotes were dissociated anteroposteriorly in a colubrid snake, these observations support the hypothesis that structures usually associated with the neck-trunk boundary in quadrupedal squamates are displaced relative to one another in snakes. Whereas certain craniovertebral muscles are elongated in some snakes, results of optimization on an ophidian cladogram show that the most recent common ancestor of extant snakes would have had the longest craniovertebral muscle, M. rectus capitis anterior, that is elongated only by several segments compared with that of quadrupedal squamates. Therefore, even such a posteriorly displaced "cervical" characteristic plesiomorphically lies fairly anteriorly in the greatly elongated precloacal region of snakes, suggesting that the trunk, not the neck, would have contributed most to the elongation of the snake precloacal region. A similar dissociation of structures usually associated with the neck-trunk boundary in quadrupedal squamates is observed in limb-reduced squamates, suggesting that these forms and snakes may share a developmental mechanism producing modifications in the anterior-posterior patterning associated with body elongation.