Osteohistology of the hyperelongate hemispinous processes of Amargasaurus cazaui (Dinosauria: Sauropoda): Implications for soft tissue reconstruction and functional significance

Abstract

Dicraeosaurid sauropods are iconically characterized by the presence of elongate hemispinous processes in presacral vertebrae. These hemispinous processes can show an extreme degree of elongation, such as in the Argentinean forms Amargasaurus cazaui, Pilmatueia faundezi and Bajadasaurus pronuspinax. These hyperelongated hemispinous processes have been variably interpreted as a support structure for a padded crest/sail as a display, a bison-like hump or as the internal osseous cores of cervical horns. With the purpose to test these hypotheses, here we analyze, for the first time, the external morphology, internal microanatomy and bone microstructure of the hemispinous processes from the holotype of Amargasaurus, in addition to a second dicraeosaurid indet. (also from the La Amarga Formatin; Lower Cretaceous, Argentina). Transverse thin-sections sampled from the proximal, mid and distal portions of both cervical and dorsal hemispinous processes reveal that the cortical bone is formed by highly vascularized fibrolamellar bone interrupted with cyclical growth marks. Obliquely oriented Sharpey's fibres are mostly located in the medial and lateral portions of the cortex. Secondary remodelling is evidenced by the presence of abundant secondary osteons irregularly distributed within the cortex. Both anatomical and histological evidence does not support the presence of a keratinized sheath (i.e. horn) covering the hyperelongated hemispinous processes of Amargasaurus, and either, using a parsimonious criterium, in other dicraeosaurids with similar vertebral morphology. The spatial distribution and relative orientation of the Sharpey's fibres suggest the presence of an important system of interspinous ligaments that possibly connect successive hemispinous processes in Amargasaurus. These ligaments were distributed along the entirety of the hemispinous processes. The differential distribution of secondary osteons indicates that the cervical hemispinous processes of Amargasaurus were subjected to mechanical forces that generated higher compression strain on the anterior side of the elements. Current data support the hypothesis for the presence of a ‘cervical sail’ in Amargasaurus and other dicraeosaurids.