An Investigation Into Ichthyosaur Ontogeny, Sexual Dimorphism and Body Size Evolution
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AN INVESTIGATION INTO ICHTHYOSAUR ONTOGENY, SEXUAL DIMORPHISM AND BODY SIZE EVOLUTION SAMUEL PATRICK BENNETT Thesis submitted for the degree of Doctor of Philosophy Royal Holloway College 1 Declaration of Authorship I, Samuel Patrick Bennett, hereby declare that this thesis and the work presented in it is entirely my own. Where I have consulted the work of others, this is always clearly stated Signed: Date: 2 ABSTRACT Ichthyosaurs are an extinct group of diapsid marine reptiles that existed from the Olenekian (251Ma) to end Cenomanian (93.9Ma). Morphometric data (length measurements) and meristic data (counts of repeated elements) were collected for Lower and Middle Jurassic taxa from several museums in England and one in Germany. Additional morphometric data were collected from the published record. Principal Components Analysis (PCA), Principal Coordinate Analysis (PCO) and Reduced Major Axis Regression (RMA) were used to analyse morphometric data relating to ontogeny. Linear Regression, also known as Ordinary Least Squares (OLS) was used to analyse meristic data relating to ontogeny as well as body size evolution. Sexual dimorphism was analysed using the Mann-Whitney test as well as Discriminant Analysis. The analysis of ichthyosaur ontogeny showed that neonate and juvenile ichthyosaurs had significantly larger skulls and eyes compared to body length. Once maturity is reached growth becomes isometric, and no other features varied with relative age. The numbers of repeated elements in ichthyosaurs remain stable throughout life, with the exception of post- flexural caudal vertebrae, where the numbers increase with the size of the tail, and therefore, age. Investigation of sexual dimorphism indicated that inferred males are isometrically larger than pregnant females. However, the age at which maturity is reached cannot currently be identified in individual specimens. Furthermore, the gender of an individual cannot currently be determined, with the exception of pregnant females. The study of body size evolution was not conclusive and no statistically significant trends were identified. Due to the nature of the fossil record, only 53% of the taxa examined could be used in the analysis. More taxa, particularly from the Triassic and Cretaceous, need to be included to improve the understanding of ichthyosaur body size evolution. 3 CONTENTS Abstract . 3 Contents . 4 Contents: Figures . 13 Contents: Tables . 19 Acknowledgements . 20 1: Ichthyosaur ontogeny, sexual dimorphism and body size evolution: an introduction . 21 1.1: Introduction to ichthyosaurs . 21 1.2: Introduction to ontogeny . 24 1.3: Introduction to sexual dimorphism . 25 1.4: Introduction to body size evolution . 26 1.5: Aims . 27 1.6: Thesis overview . 28 2: Methodology . 30 2.1: Introduction . 30 2.1.1: Overview . 30 2.2: Materials . .31 2.2.1: Institutions . 31 2.2.2: Genera . 31 2.2.3: Taxonomy of the Lower Jurassic ichthyosaur taxa used in this study .32 2.2.4: Key implications for the thesis . 59 4 2.3: Morphometric data . 59 2.4: Meristic (counted) data . 61 2.5 Multivariate analysis . 62 2.5.1: Principal Components Analysis (PCA) . 62 2.5.2: Principal Coordinate Analysis (PCO) . 66 2.5.3: Reduced Major Axis Regression (RMA) . 67 2.5.3.1: Bootstrap test . 68 2.5.3.2: Pearson’s R Correlation . 69 2.5.3.3: Discriminant Analysis . 69 3: Investigating ichthyosaur growth: a morphometric approach . 74 3.1: Overview . 74 3.2: Definitions: ontogeny, heterochrony and allometry . 75 3.3: The importance of understanding ontogeny . 77 3.4: Approaches to studying diapsid ontogeny in the fossil record . 78 3.4.1: An introduction to ichthyosaur ontogeny . .79 3.4.2: Size independent criteria . 80 3.4.3: Bone histology . 82 3.4.4: Tooth loss . 85 3.4.5: Ossification patterns . 86 3.4.6: Other ontogenetic differences . 88 3.4.7: Discussion of ichthyosaur ontogeny . 88 3.4.8: Summary of current knowledge of ichthyosaur ontogeny . 90 5 3.5: Ontogeny of other diapsids: Sauropterygia . 64 3.5.1: Plesiosaurs . 91 3.5.2: Pachypleurosaurs . 92 3.5.2.1: Inferred embryos . 93 3.5.2.2: Inferred juveniles . 94 3.5.2.3: Inferred adults . 95 3.5.2.4: Bone histology . 95 3.5.2.5: Sexual dimorphism in pachypleurosaurs . 96 3.5.2.6: Pachypleurosaur summary . 96 3.6: Ontogeny of other diapsids: Crocodylia . 97 3.6.1: Bone fusion . 97 3.6.2: Other ontogenetic features . 98 3.7: Ontogeny of other diapsids: Dinosauria . 99 3.7.1: Triceratops . 99 3.8: Summary of evidence for ontogeny . 100 3.9: Implications for ichthyosaur ontogeny . 101 3.10: Materials and methods . 101 3.11: Results and interpretation . 102 3.11.1: Multivariate analysis: PCA . 102 3.11.1.1: PCA analysis: humerus . 103 3.11.1.2: PCA Analysis: femur . 108 3.11.1.3: PCA Analysis: skull . 113 3.11.1.4: PCA analysis of all measurements . 118 6 3.11.2: Multivariate analysis: PCO . 119 3.11.2.1: PCO analysis: humerus . 119 3.11.2.2: PCO Analysis: femur . 120 3.11.2.3: PCO Analysis: skull . 122 3.11.3: Bivariate analysis: RMA . 123 3.11.3.1: All measurements vs. total body length . 124 3.11.3.2: Humerus length vs. humerus width . 131 3.11.3.3: Femur length vs. femur width . 132 3.11.3.4: Skull length vs. jaw length . 133 3.11.3.5: Length of longest digit vs. width of paddle . 134 3.11.3.6: Total body length vs. skull length . 135 3.11.3.7: Skull length vs. external diameter of the sclerotic ring . 136 3.12: Discussion . 137 3.12.1: Limitations of the data . 137 3.12.2: Significance of results . ..