Modularity and integration of the neural arch and vertebral centrum in primates

The vertebral column consists of multiple homologous elements that have specialized within and between taxa and serve important functions in positional support and as protection for the central nervous system. The study of modularity and integration provides new insights into the evolution of complex structures such as the vertebral column. Patterns of modularity and integration may reflect underlying genetic-developmental patterns and facilitate evolution. Previous studies have identified mixed modularity patterns within and between elements across mammals generally, within primates and carnivorans. Here, we assess modularity within and between elements in the complete post-axial vertebral column in four catarrhine taxa: Macaca (n = 96), Hylobates (n = 77), Pan (n = 92), and Homo (n = 151). We use the Covariance Ratio (CR) to estimate r² and the standardized eigenvalues (SVE) variance for comparative purposes. Our results show that there is general, widespread integration within the catarrhine vertebral column, both within and between elements. Hominoids tend to display greater modularity than do macaques, but these estimates are rarely significant. Clusters of modularity in the mid-cervical and upper thoracic regions may relate to special nervous system structures in these areas, and locomotor behaviors in general may influence patterns of modularity in primates. In particular, we find that size is a pervasive factor affecting integration among vertebral elements, though its effects on specific structures are variable. Our results generally do not agree with those found across mammals or within carnivorans, and future studies should focus on genus-level assessments across a variety of taxa.