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Osteohistology and Growth of Edmontosaurus and Assessment of Life History Changes in Hadrosaurid Dinosaurs

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Osteohistology and Growth of Edmontosaurus and Assessment of Life History Changes in Hadrosaurid Dinosaurs Osteohistology and Growth of Edmontosaurus and Assessment of Life History Changes in Hadrosaurid Dinosaurs by Mateusz Wosik A thesis submitted in conformity with the requirements for the degree of Doctor of Philosophy Department of Ecology and Evolutionary Biology University of Toronto © Copyright by Mateusz Wosik 2018 Osteohistology and Growth of Edmontosaurus and Assessment of Life History Changes in Hadrosaurid Dinosaurs Mateusz Wosik Doctor of Philosophy Department of Ecology and Evolutionary Biology University of Toronto 2018 Abstract Data on the life history of an organism provides an important window into its biology, ecology, and evolution but has been challenging to obtain for the extinct due to limitations of fossil sampling and the inability for direct observation. The recent utilization of osteohistology, which is the study of the internal bone microstructure, has revolutionized paleontological research and permitted quantitative assessment of numerous life history aspects in extinct organisms. Hadrosaurid dinosaurs, the dominant large-bodied terrestrial herbivores in most Late Cretaceous ecosystems, have an exceptional fossil record consisting of over 60 species and multiple ontogenetic bonebeds making them an ideal clade on which to conduct life history studies. The objectives of this thesis were to integrate new data and methods to investigate ontogenetic growth and test life history strategies in hadrosaurid populations from the Late Cretaceous of North America, with a particular emphasis on the integration of osteohistological methods with limb growth and size frequency distributions for inferring biomechanical gait shifts, growth and population dynamics, and mass death preservational patterns. The results in this thesis demonstrate that the proportions of the appendicular skeleton of hadrosaurid dinosaurs did not undergo allometric growth through ontogeny and provide no compelling evidence to support the ii previously hypothesized ontogenetic gait shift in hadrosaurids. Size-frequency distributions underestimate age and overestimate growth rates. However, when paired with osteohistology, the data reveal that juvenile hadrosaurids segregated from adults providing essential insight into the population dynamics of these animals. The comprehensive growth curve analysis indicates that hadrosaurids exhibited a similar growth trajectory regardless of a wide ranging asymptotic body size (~3000–6000 kg) suggesting that the clade as a whole inherited a similar growth strategy attaining adult size within 7–9 years. This thesis strongly recommends both the independent assessment and the integration of allometric scaling, size-frequency distributions, and osteohistology to provide more accurate and precise paleobiological interpretations of life history characteristics. iii © Dennis M. Yeider A Day in the Life of Edmontosaurus. Edmontosaurus nestlings, similar to many other duck- billed dinosaurs, segregated from adults likely forming their own independent groups. Here, one of these groups encounters the skeletal remains of an adult. The illustration of the Sandstone Basin locality from northeastern Montana is split into a present field photograph (left) and interpretation of the paleoenvironment (right) to demonstrate the significant environmental changes over 66 million years. iv I dedicate this thesis to my son, Otter Emmerich Wosik. Your first year of life has known nothing but me working on this thesis. With it behind us, know that I will always be there for you. Now let’s enjoy the adventures our lives have in store for us! v Acknowledgments So many people have helped me along the way during this work that I sincerely hope that I do not forget anyone. To start, I’d like to thank: …My advisor and mentor, Dr. David C. Evans. When I entered the lab, I was the definition of green. Full of wild ideas, no focus, and chasing rabbit holes. If you must know, I did indeed read large portions of that Zar biostats book but still have no idea how to reasonably comprehend quantile-quantile plots. During my PhD, you molded my mind to be more than just a scientist by providing me with a wide array of opportunities to develop to various skillsets. My favorite was when you let me literally shovel dirt for hours, and on more than one occasion (true fact, I love shoveling me some dirt). But more seriously, you taught me the patience to fully analyze each situation and the perseverance of commitment. You were supportive when life took unexpected detours, and at times, felt isolating. I know I wasn’t the easiest of students and occasionally tested your nerves, but I truly hope that one day, I can follow your lead and pass on the wisdom you shared with me. …My supervisory committee members, Dr. Deborah McLennan and Dr. Robert Reisz, for making me think beyond the scope of this work by relating it to the modern world. Your constructive feedback, and the occasional complete rewrite of sections, significantly shaped this work from its infancy. An honorable mention to the science related comic strips that provided much needed self-esteem boosts when data and results had a mind of their own. …My appraisal and defense committee members, Dr. Anusuya Chinsamy-Turan, Dr. Marc LaFlamme, Dr. Mary Silcox, and Dr. Bob Murphy for critically testing my current knowledge and stretching it beyond lengths I could have never imagined. vi …My undergraduate professors that steered me towards zoology and ecology, in particular Dr. Kyle Bennett, Dr. Chris Petersen, Dr. Paul Arriola, and especially Dr. Merrilee Guenther, who offered me my first opportunity in paleontology as an undergraduate research assistant and single handedly saved me from the dark side of molecular biology. …The ROM paleo staff, Peter Fenton, Brian Iwama, Ian Morrison, Dave Rudkin, Kevin Seymour, and Shino Sugimoto, I cannot thank you enough. Beyond all the time that you invested to teach me new skills, I want you to know that your continued support and words of encouragement did not go unnoticed, and at times, had more of a positive impact than I let up. …All of my ROM labmates, past and present, with special mention to Kentaro Chiba, Aaron Leblanc, Cary Woodruff, and Danielle Dufault for having the patience to answer all my questions, teaching me all I know about histology, and being an ear when I needed to get things off my chest. From noodles to beers, I’ll always cherish the memories of growing up with you. …The entire Badlands National Park staff for being at the core of the two best summers of my life. I arrived in a mysterious land not knowing a single soul, but left with a heavy heart and countless life-long relationships. To the staff and volunteers that taught me all I know about outreach: Rachel Benton, Megan Cherry, Connie Wolf, Paul, Larry, Cindy, Chuck, and Sandra. To the friends that made every laugh count: Levi Darwin Moxness, Dakota McCoy, Ed Welsh, Nick Famoso, Sarah Nevison, Danny Redding, Evan Doughty, Ian Knoerl, Clint Boyd, Mindy Householder, Katie Johnston, and so many more. And most importantly, Aaron Kaye and Ellen Starck. Without your encouragement and support, I may have never considered continuing my education. For that alone, have a proper liquid courage and a chilled Mountain Dew, on me. vii I am also very grateful for the financial support during my time at the University of Toronto. A five year University of Toronto Fellowship, four one-year Joseph Bazylewicz Fellowships, as well as numerous departmental-level awards and travel funds made this work possible. To my dear friends: Sungsik ‘Kevin’ Kong, Tomas Frecka, Eva Rodinova, Peter Vlcek, Magdalena Salatova, Laurel Duquette, Eric Owens, Mike Swift, and especially Sarah Steele. I am going to bloody damn miss you all. From countless laughs to tearful goodbyes, you made living in another country that much easier because I knew I could rely on each of you for anything. Although we may be friends by blood, you all are my family by heart. To my in-laws, Dennis and Cynthia Yeider. I will always remember that before I could finish asking you for your permission to marry your daughter, you interjected to say ‘absolutely yes.’ From that moment, I knew I was being welcomed into an incredible family that has shown me nothing but endless support ever since, especially during my Ph.D. studies. To my dog, Moose. Thank you for putting up with my near-criminal practice talks and keeping me in check by giving your infamous cue when it was time to go to bed. To my wife, Lindsey Elizabeth Yeider, I thank you for your endless love and support. Through stratospheric highs and earth shattering lows, you have been the single constant in my life. Whenever I needed a bit of a nudge, you jumped right into the water with me. If I ever felt knocked down, you dug a 10 foot hole to show me I was nowhere near the bottom. I am truly blessed knowing that I married my best friend. When we moved to Toronto nearly five years ago, I never imagined that we would be concluding this chapter of our lives as a family. Let’s never forget that ‘adventure is out there’ and now we have a ‘zoo’ to tag along with us. viii And lastly, to my parents, Maria and Stanisław Wosik. Przede wszystkim nauczyliście mnie wartośći ciężkiej pracy i dopilnowywania, bym kończył to, co zacząłem. Chociaż nigdy nie będę w stanie odwdzięczyć się za Wasze bezinteresowne poświęcenie w czasie, kiedy wyemigrowaliśmy do Stanów Zjednoczonych, zapewniliście mi tą możliwość, która mam nadzieję, czyni was dumnymi. Z głebi serca, dziękuję Wam serdecznie. Above all, you taught me the value of hard work and making sure to finish what I started. Although I will never be able to repay the selfless sacrifices the both of you endured when we immigrated to the United States, you provided me with this opportunity that I hope makes you proud.
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