Morphological variation of the thoracolumbar vertebrae in Macropodidae and its functional relevance.
The present study was designed to investigate how the form of the marsupial thoracolumbar vertebrae varies to cope with the particular demands of diverse loading and locomotor behaviors. The vertebral columns of 10 species of Macropodidae, with various body masses and modes of locomotion, together with two other arboreal marsupials, koala and cuscus, were selected. Seventy-four three-dimensional landmark coordinates were acquired on each of the 10 last presacral vertebrae of the 70 vertebral columns. The interspecific variations of the third lumbar vertebra (L3, which approximates the mean) and the transitional patterns of the thoracolumbar segments were examined using the combined approach of generalized Procrustes analysis (GPA) and principal components analysis (PCA). The results of analyses of an individual vertebra (L3) and of the transitional patterns indicate significant interspecific differences. In the L3 study the first PC shows allometric shape variation, while the second PC seems to relate to adaptation for terrestrial versus arboreal locomotion. When the L3 vertebrae of the common spotted cuscus and koala are included for comparison, the vertebra of the tree kangaroo occupies an intermediate position between the hopping kangaroo and these arboreal marsupials. The L3 vertebrae in the arboreal marsupials possess a distinct dorsoventrally expanded vertebral body, and perpendicularly orientated spinous and transverse processes. The results of the present study suggest that vertebral shape in the kangaroo and wallaroos provides a structural adaptation to hopping through a relatively enlarged loading area and powerful lever system. In contrast, the small-sized bettongs (or rat kangaroos) have a relatively flexible column and elongated levers for the action of back muscles that extend and laterally flex the spine. The complex pattern of vertebral shape transition in the last 10 presacral vertebrae was examined using PCAs that compare between species information about vertebral shape variation along the thoracolumbar column. The results reinforce and emphasize important aspects of the patterns of variation seen in the detailed analysis of the third lumbar vertebra. The results also imply that size, spinal loading pattern, and locomotor behavior exert an influence on shaping the vertebra. Further, the morphological adaptations are consistent within these marsupials and this opens up the possibility that this kind of analysis may be useful in making functional inferences from fossil material.