Osteohistology of the unusually fast-growing theropod dinosaur Ceratosaurus.

Ceratosaurus is a large-bodied non-avian theropod dinosaur known from the Upper Jurassic Morrison Formation of North America and is remarkable both for its exceptionally fast annual growth rate and its status as the only theropod currently known with postcranial osteoderms. We describe the osteohistology of three hind limb bones, two dorsal ribs, and one osteoderm representing four individuals of Ceratosaurus. In addition to describing the tissues of these bones, we compared the annual growth rates from three individuals in our sample to those of five other ceratosaurians. We fit seven growth models to two of the specimens in our sample and compared the results of the best-fit model(s) to those of two other ceratosaurians (Masiakasaurus knopfleri and Majungasaurus crenatissimus) for which sufficient growth data were available. The bone tissue of hind limbs in Ceratosaurus is highly vascularized, with dense plexiform or reticular vascular complexes and alternating strips of parallel or woven-fibered matrix. Few lines of arrested growth were recorded in hind limbs prior to specimens achieving asymptotic body size. Both sampled dorsal ribs are highly remodeled, with only small portions of primary bone visible in each section, revealing parallel-fibered bone with sparse primary osteons. Both dorsal ribs contain numerous lines of arrested growth throughout the cortex that allowed for more accurate estimates of individual age when paired with the data from hind limbs. The osteoderm is composed of a core of large Haversian canals and a perimeter of lamellar bone with dense Sharpey's fibers along the internal surface of the bone. Multiple LAGs are also present within the lamellar bone along the exterior margins. Maximum annual growth rates in Ceratosaurus were on average nine-fold faster than those of other ceratosaurians. Our sample lacks data from juveniles so confidence in inferred growth models is limited. Thus, to begin to constrain Ceratosaurus growth patterns, we averaged the results of all models that possessed an Akaike Information Criterion score corrected for small sample size (AICc) within 10 of the lowest scoring model. We found that the monomolecular model exhibited the lowest AICc value, with the von Bertalanffy and Gompertz models possessing AICc values within 10 units of it. In contrast, the logistic and Gompertz models were confidently selected for Masiakasaurus and Majungasaurus, respectively. Irrespective of growth model, maximum relative annual growth rates for Ceratosaurus were several-fold greater than those of Masiakasaurus and Majungasaurus. Both histological and growth model estimates of life history support an evolutionary trend towards more prolonged development in Ceratosauria through evolutionary time.