Evolution of the vertebral formulae in mammals: a perspective on developmental constraints.

Developmental constraints refer to biases that limit phenotypic changes during evolution. To examine the contribution of developmental constraints in the evolution of vertebrate morphology, we analyzed the distribution pattern of mammalian vertebral formulae. Data on mammalian vertebral formulae were collected from the Descriptive Catalogue of the Osteological Series Contained in the Museum of the Royal College of Surgeons of England by Richard Owen (1853) and were plotted onto the most reliable mammalian phylogenetic tree based on recent molecular studies. In addition to the number of cervical vertebrae that is almost fixed to 7, we found that the number of thoracolumbar vertebrae tends to be 19 in many groups of mammals. Since fidelity of the number of thoracolumbar vertebrae was also completely maintained in Monotremata and Marsupialia, we presumed that thoracolumbar vertebral number as well as cervical vertebral number might have been fixed in the primitive mammalian lineage. On the basis of primitive vertebral formulae, we could clarify the polarity of evolution and identify several deviations from the primitive states during the mammalian evolution. The changes in the vertebral formulae in eutherian mammals seem to be lineage-specific, such that most species in Carnivora have 20 instead of 19 thoracolumbar vertebrae. Because such lineage-specific vertebral formulae contrast with the estimated distribution pattern on the assumption of evolution only through the selective pressure, we concluded that developmental constraints played an important role in the evolution of mammalian vertebral formulae.