Paleobiology of ichthyosaurs: using osteohistology to test hypotheses of growth rates and metabolism in a clade of secondarily aquatic marine tetrapods

Abstract

Ichthyosaurians (Ichthyosauria) are one of the most prominent groups of secondarily aquatic Mesozoic marine reptiles. Over their 160 million years of evolution, the clade evolved a streamlined body plan with paddle-like limbs, convergent with modern cetaceans. Despite the fact that ichthyosaurians have been studied by paleontologists for over a century, very little is known about aspects of their biology, including quantification of their age structure and growth rates. Multiple lines of evidence, including oxygen isotope, swimming modality, and body shape analyses suggest that ichthyosaurians experienced elevated growth rates and likely maintained an elevated body temperature relative to ambient sea water. In this dissertation, I test these hypotheses using osteohistological methods. In the first manuscript, we describe new material of the small-bodied Upper Triassic ichthyosaurian Toretocnemus from the Nehenta Formation and the Hound Island Volcanics (both Norian, Upper Triassic) of Southeast Alaska. During the Upper Triassic, ichthyosaurians experienced their greatest size disparity, with large-bodied species rivaling the size of modern blue whales (Balaenoptera musculus; 20+ m body length) living alongside small-bodied species (1 m body length) like Toretocnemus. Prior to this study, Toretocnemus was known from Carnian deposits of California and possibly Sonora, Mexico. The referred material described here expands its geographic and temporal range. There are very few known ichthyosaurians from the Norian; thus, this material sheds light on the clade's diversity before the end Triassic extinction event. In the second and third manuscripts, we use osteohistological methods to describe the microstructure of various skeletal elements of two species of Stenopterygius from the Posidonia Shale (Lower Jurassic) of Germany. The Posidonia Shale is a Konservat-Lagerstätten that preserves over 3000 ichthyosaurian specimens, approximately 80 percent of which are referable to Stenopterygius. First, we sampled over 40 skeletal elements from one individual specimen referred to Stenopterygius quadriscissus to 1) describe the mineralized tissues across the skeleton, 2) infer relative growth rate, and 3) identify elements with growth marks. Almost all elements described demonstrate fibrolamellar primary bone, indicative of a rapid growth rate. We also identify growth marks in several elements, including the dentary and premaxilla, that will be used in future growth studies. In the third manuscript, we sample a scleral ossicle from Stenopterygius triscissus to describe its microstructure and investigate the use of ossicles for skeletochronology. The use of scleral ossicles for determining age structure has been documented in extant sea turtles as well as dinosaurs. We sectioned one ossicle in three planes and document conspicuous growth banding in the short axis section. Although this method requires further testing, we tentatively determine a minimum age of 7 years at the time of death for this individual. This dissertation lays critical groundwork for future studies of the paleobiology of ichthyosaurians. We are already in the preliminary stages of using these results to 1) quantify age structure and growth rates of an ichthyosaurian (Stenopterygius quadriscissus) for the first time, and 2) test the use of scleral ossicles for skeletochronology of ichthyosaurians. Through addressing these basic aspects of ichthyosaurian biology, we can begin to investigate how ichthyosaurian development and physiology changed over time and space and develop a greater understanding of this clade's 160 million years of evolution.