Interelemental osteohistological variation in Massospondylus carinatus and its implications for locomotion

Massospondylus carinatus Owen, 1854 is an iconic basal sauropodomorph dinosaur from the Early Jurassic of southern Africa. Over 200 specimens have been referred to this taxon, spanning the entire ontogenetic series from embryo to adult. Consequently, it provides an ideal sample for investigating dinosaur developmental biology, including growth patterns and growth rates, through osteohistological analysis. Massospondylus carinatus was the first early-branching sauropodomorph dinosaur for which a femoral growth series was sampled. Since then, growth series of other non-avian dinosaur taxa have shown that growth plasticity, interelemental variation, and ontogenetic locomotory shifts can complicate our understanding of growth curves and patterns. To investigate these questions further, it is necessary to sample multiple skeletal elements from multiple individuals across a large range of sizes, something that is often hindered by the incompleteness of the fossil record. Here, we conducted a broad, multielement osteohistological study of long bones (excluding metapodials) from 27 specimens of Massospondylus carinatus that span its ontogenetic series. Our study reveals substantial variations in growth history. A cyclical woven-parallel complex is the predominant bone tissue pattern during early and mid-ontogeny, which transitions to slower forming parallel-fibred bone during very late ontogeny. The bone tissue is interrupted by irregularly spaced cyclical growth marks (CGMs) including lines of arrested growth indicating temporary cessations in growth. These CGMs show that the previously recorded femoral growth plasticity is also visible in other long bones, with a poor correlation between body size (measured by midshaft circumference) and CGM numbers. Furthermore, we found that the growth trajectory for an individual can vary depending on which limb element is studied. This makes the establishment of an accurate growth curve and determination of the onset of reproductive maturity difficult for this taxon. Finally, we found no evidence of differential growth rates in forelimb vs hindlimb samples from the same individual, providing further evidence falsifying hypothesised ontogenetic postural shifts in Massospondylus carinatus.