Evolutionary Dynamics of Tyrannosauroid Dinosaurs

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Evolutionary Dynamics of Tyrannosauroid Dinosaurs Facultat de Ciències Memòria del Treball Final de Grau Títol del treball: Evolutionary Dynamics of Tyrannosauroid Dinosaurs Estudiant: Laia Garcia Escolà Grau en biologia Correu electrònic: [email protected] Tutor: Sandra Heras Mena Cotutor: Albert Prieto Márquez Empresa/institució: ICP (Institut Català de Paleontologia) Vistiplau tutor (i cotutor): Nom del tutor: Sandra Heras Mena ([email protected]) Nom del cotutor: Albert Prieto Márquez ([email protected]) Empresa/ institució: Institut Català de Paleontologia Miquel Crusafont Data de dipòsit de la memòria a secretaria de coordinació: 03/06/2019 Evolutionary Dynamics of Tyrannosauroid Dinosaurs Laia Garcia Escolà Abstract This study explores the evolutionary dynamics of tyrannosauroid dinosaurs, a successful clade of basal coelurosaur theropods. This is accomplished by estimating rates of evolution using the Claddis R package. Morphological diversity was measured from discrete character data and estimates of evolutionary tempo on current phylogenetic hypotheses of tyrannosauroid relationships. In this work it is shown the process to obtain the set of necessary documents to obtain the results with the script to finally calculated it with R. In this set of previous documents are included a matrix character, that gather a series of different characters associated with the different taxa into the clade. It is also added a temporary range of each taxon and a consensus phylogenetic tree made by phylogenetic analysis under parsimony. All of them obtained through different software (Mesquite, TNT and R) and necessaries to run the script to obtain the seek resultants rates. Evolutionary rates were estimated for different anatomical regions of the tyrannosauroid skeleton, including the skull and mandible, the post cranial body and the whole-body. These phylogenetic trees include the rate calculation, the mode and tempo of changes produced in each node, to perceive possible acceleration or deacceleration of morphological evolution on the different branches of this clade. To help with the conclusions and discussions of the study it is assessed a relationship between tyrannosauroid evolution and the body mass of their potential prey, ceratopsians and ornithopods through time. The results indicate that the mandible was a driver of the increasing mass of the skull in tyrannosauroids, accompanied by an increase of the size of the posterior region of the skeleton and the reduction of the forelimbs. An important conclusion of this study is that the increasing skull size facilitates a greater bite force. The relationship between tyrannosauroids body mass and that of their potential preys indicates that is boosting, which could be a case of arms race, being a potential cause of this mandible adaptation, which is supported by different studies. I Evolutionary Dynamics of Tyrannosauroid Dinosaurs Laia Garcia Escolà Resum Aquest treball estudia la dinàmica evolutiva dels dinosaures tiranosauroides, un exitós clade basal de teròpodes coelurosaures. Això s’aconsegueix mitjançant l'estimació de les taxes d’evolució utilitzant un paquet estadístic, anomenat Claddis, per al programa R. La diversitat morfològica es mesura a partir de dades de caràcter discret i estimacions del temps evolutiu sobre les hipòtesis filogenètiques actuals de les relacions dels tiranosauroides. En aquests treball es mostra el procés per a obtenir un conjunt de documents necessaris per a poder obtenir els resultats a través d’un script i que puguin ser finalment calculats amb el programa estadístic R. En aquest conjunt de documents anteriors s’inclouen una matriu de caràcters, que recull una sèrie de diferents caràcters associats als diferents tàxons del clade. També s’afegeix un rang temporal de cada taxó i un arbre filogenètic consens obtingut a partir d’un anàlisi filogenètic sota parsimònia. Tots ells, necessaris per executar l’script, obtenir les taxes resultants i obtinguts a través de diferents programes (Mesquite, TNT i R). Les taxes evolutives es calculen per a cada region anatòmica de l’esquelet dels tiranosauroides, les quals es divideixen en el crani i la mandíbula, el crani sense la mandíbula, en el cos post cranial i en el cos sencer. Aquests arbres filogenètics inclouen el càlcul de la velocitat, el mode i el temps dels canvis produïts a cada node, per percebre una possible acceleració o desacceleració de l’evolució morfològica a les diferents branques d’aquest clade. Per ajudar amb les conclusions i la discussió sobre l’estudi, s’avalua la relació entre l’evolució de la massa corporal dels tiranosauroides i de les seves preses potencials, ceratopsians i ornitòpodes a través del temps. Finalment, els resultats indiquen que la mandíbula va ser un dels principals motors en l’augment de la massa del crani en tiranosauroides, acompanyada d’un augment de la mida de la regió posterior de l’esquelet i la reducció de les extremitats anteriors. Una conclusió important d’aquest estudi és que l’augment de la mida del crani facilita una major força de mossegada. La relació entre la massa corporal dels tiranosauroides i la de les seves preses potencials indica que està augmentant de manera similar, cosa que podria ser un cas d’ “arms race”, sent la causa potencial d'aquesta adaptació de la mandíbula i recolzada en diferents estudis. II Evolutionary Dynamics of Tyrannosauroid Dinosaurs Laia Garcia Escolà Resumen Este trabajo estudia la dinámica evolutiva de los dinosaurios tiranosauroides, un exitoso clado basal de terópodos coelurosaures. Esto se consigue mediante la estimación de las tasas de evolución utilizando el paquete estadístico Claddis para R. La diversidad morfológica se mide a partir de datos de carácter discreto y estimaciones del tiempo evolutivo sobre las hipótesis filogenéticas actuales de las relaciones de los tiranosauroides. En este trabajo se muestra el proceso de obtención de un conjunto de documentos necesarios para el calculo mediante un script para R. En este conjunto de documentos anteriores incluyen una matriz de caracteres, que recoge una serie de diferentes caracteres asociados a los diferentes taxones del clado. También se añade un rango temporal de cada taxón y un árbol filogenético consenso obtenido a partir de un análisis filogenético bajo parsimonia. Todos ellos, necesarios para ejecutar el script, obtener las tasas resultantes y obtenidos a través de diferentes programas (Mesquite, TNT y R). Las tasas evolutivas se calculan para cada región anatómica del esqueleto de los tiranosauroides, las cuales se dividen en el cráneo y la mandíbula, el cráneo sin la mandíbula, en el cuerpo post craneal y en el cuerpo entero. Estos árboles filogenéticos incluyen el cálculo de la velocidad, el modo y el tiempo de los cambios producidos en cada nodo, para percibir una posible aceleración o desaceleración de la evolución morfológica en las diferentes ramas de esta categoría filogenética. Para ayudar con las conclusiones y la discusión sobre el estudio, se evalúa la relación entre la evolución de la masa corporal de los tiranosauroides y de sus presas potenciales, ceratopsians y ornitópodos a través del tiempo. Finalmente, los resultados indican que la mandíbula fue uno de los principales motores en el aumento de la masa del cráneo en tiranosauroides, acompañada de un aumento del tamaño de la región posterior del esqueleto y la reducción de las extremidades anteriores. Una conclusión importante de este estudio es que el aumento del tamaño del cráneo facilita una mayor fuerza de mordida. La relación entre la masa corporal de los tiranosauroides y la de sus presas potenciales indica que está aumentando de manera similar, lo que podría ser un caso de “arms race”, siendo la causa potencial de esta adaptación de la mandíbula y apoyada en diferentes estudios. III Index Introduction ………….………………………..……….………..…..…… 2 Objectives ……………………………………………….……….…….…… 5 Methodology …………………………………………………………..…. 6 Ethics and sustainability …………..…………………………….… 11 Results ……………………………………………………………………… 11 Discussion ……………………………………………….………………… 20 Conclusion …………………………………………..……………………. 22 Acknowledgment …………………..…………………………….…… 22 References ………………………………………….…………………….. 23 Evolutionary Dynamics of Tyrannosauroid Dinosaurs Laia Garcia Escolà Introduction Tyrannosauroids, are a clade of theropod dinosaurs spanning the Middle Jurassic through the latest Cretaceous (Osborn, 1905; Gauthier, 1986; Brussate et al., 2010). Their fossil record is distributed among Eurasia, the Americas and Australia (Lyson, 2010) (Figure 1). Although is been revealed in recent studies that the distribution of tyrannosauroids could have been cosmopolitan it’s not formally probed, as some authors believed that tyrannosauroids were absent from the most arid paleoenvironments (Holtz, 2004). Figure 1. Representation map of the tyrannosauroids distribution in an actual Earth map. Colours on the map match with the colours of the geological periods below. This clade includes some of the most iconic carnivorous dinosaurs, such as Tyrannosaurus rex. Tyrannosauroids have been an iconic clade since the beginning of the twentieth century. (Osborn, 1905). For much of the century, these animals were included within the carnosaur group (Brussate et al., 2010). Carnosaurs included all the large theropods, some of them of gigantic size, like T. rex. 2 Evolutionary Dynamics of Tyrannosauroid Dinosaurs Laia Garcia Escolà Before the advent of cladistics, the relationship of dinosaurs and other fossil organisms followed a Linnaean classification scheme. Subsequently,
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