Morphology, Phylogeny, and Evolution of Diadectidae (Cotylosauria: Diadectomorpha)
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Morphology, Phylogeny, and Evolution of Diadectidae (Cotylosauria: Diadectomorpha) by Richard Kissel A thesis submitted in conformity with the requirements for the degree of doctor of philosophy Graduate Department of Ecology & Evolutionary Biology University of Toronto © Copyright by Richard Kissel 2010 Morphology, Phylogeny, and Evolution of Diadectidae (Cotylosauria: Diadectomorpha) Richard Kissel Doctor of Philosophy Graduate Department of Ecology & Evolutionary Biology University of Toronto 2010 Abstract Based on dental, cranial, and postcranial anatomy, members of the Permo-Carboniferous clade Diadectidae are generally regarded as the earliest tetrapods capable of processing high-fiber plant material; presented here is a review of diadectid morphology, phylogeny, taxonomy, and paleozoogeography. Phylogenetic analyses support the monophyly of Diadectidae within Diadectomorpha, the sister-group to Amniota, with Limnoscelis as the sister-taxon to Tseajaia + Diadectidae. Analysis of diadectid interrelationships of all known taxa for which adequate specimens and information are known—the first of its kind conducted—positions Ambedus pusillus as the sister-taxon to all other forms, with Diadectes sanmiguelensis, Orobates pabsti, Desmatodon hesperis, Diadectes absitus, and (Diadectes sideropelicus + Diadectes tenuitectes + Diasparactus zenos) representing progressively more derived taxa in a series of nested clades. In light of these results, it is recommended herein that the species Diadectes sanmiguelensis be referred to the new genus Oradectes, Diadectes absitus be referred to the new genus Silvadectes, and Diasparactus be synonymized with Diadectes to produce Diadectes zenos. The phylogenetic hypothesis also reveals an evolutionary history leading to more efficient oral processing within the lineage, with successive nodes characterized by features indicative of a high-fiber diet. Within Diadectomorpha, diadectids constitute the majority of the species, ii suggesting that the advent of herbivory resulted in a relatively rapid radiation of species within the group, producing a clade that is markedly more species-rich than other, non-herbivorous diadectomorph taxa. An extensive review of Permo-Carboniferous tetrapod-bearing localities does, however, indicate that diadectids were not a key component of the fauna, discovered at fewer than 50 percent of the sites reviewed. These results counter suggestions that the evolution of Diadectidae led to the formation of the modern terrestrial ecosystem—where a large crop of herbivores supports a much smaller number of carnivores—during the Late Carboniferous and Early Permian. iii Acknowledgments I would like to thank my thesis committee chairperson, Dr. Robert Reisz, for his guidance, encouragement, and continued support of this project and my career. Words simply cannot express the gratitude that I will forever hold. I also wish to extend thanks to the members of my final exam committee, Drs. David Evans, Hans-Dieter Sues, Denis Walsh, and Rick Winterbottom. For her patience, teaching, and continual smile, a special thanks is offered to Ms. Diane Scott. For the loan of specimens critical to this study, I thank: Bill Amaral and Charles Schaff (Museum of Comparative Anatomy, Cambridge); Victoria Byre and Jeff Person (Sam Noble Oklahoma Museum of Natural History, Norman); Greg Gunnell (Museum of Paleontology, Ann Arbor); Brian Iwama and Kevin Seymour (Royal Ontario Museum, Toronto); Amy Henrici (Carnegie Museum of Natural History, Pittsburgh); and William Simpson (Field Museum of Natural History, Chicago). And for many discussions of diadectid anatomy, I thank Dr. David Berman of the Carnegie Museum. Finally, I thank those beautiful few that have have offered their love and support throughout not only this study but throughout my life. Thank you. iv Table of Contents Abstract ii Acknowledgments iv List of Tables viii List of Figures ix List of Appendices xv Chapter 1 Introduction and Historical Background 1 Chapter 2 Description of a New Diadected from the Lower Permian of Ohio 6 1 Introduction 6 2 Description and Comparison 8 2.1 Introduction 8 2.2 Maxilla 9 2.3 Dentary 14 2.4 Ambedus pusillus and diadectid ontogeny 18 2.5 Conclusions 19 Chapter 3 Phylogeny of Diadectidae 25 3 Introduction 25 4 Excluded Taxa 26 5 Monophyly of Diadectidae 29 5.1 Revisiting Diadectomorpha as the sister-taxon to Amniota 32 v 6 Interrelationships of Diadectidae 35 6.1 Taxonomic Implications 39 7 Conclusions 42 Chapter 4 Systematic Revision 69 8 Diadectidae Cope 1880a 69 8.1 Ambedus Kissel and Reisz 2004, p. 198 70 8.2 Orobates Berman et al. 2004, p. 3 71 8.3 Oradectes n. gen. 72 8.4 Desmatodon Case, 1908, p. 236 74 8.5 Silvadectes n. gen. 76 8.6 Diadectes Cope 1878a, p. 505 78 8.7 Problematic Taxa Assigned to Diadectidae 82 Chapter 5 Evolution of Diadectidae 91 9 Introduction 91 10 Evolutionary Radiation of Diadectidae 92 11 Evolutionary Trends within Diadectidae 94 Chapter 6 Stratigraphic and Geographic Distribution of P-C Tetrapods 99 12 Introduction 99 13 Permo-Carboniferous Tetrapod-bearing Localities 101 13.1 Missourian 102 vi 13.2 Virgilian 106 13.3 Wolfcampian 113 13.4 Leonardian 127 14 Results and Conclusions 143 References 156 Appendix 1 169 Appendix 2 173 Appendix 3 177 Appendix 4 181 vii List of Tables Table 1. Maximum crown measurements (in mm) and ratios of largest preserved, midseries maxillary cheek teeth in selected specimens of Diadectes, Desmatodon, Orobates, and Ambedus. Modified from Berman and Sumida (1995). Table 2. Maximum crown measurements (in mm) and ratios of largest preserved, midseries dentary cheek teeth in selected specimens of Diadedectes, Diasparactus, Desmatodon, Orobates, and Ambedus. Modified from Berman and Sumida (1995) and Kissel and Lehman (2002). Table 3. Missourian locality data, listing those tetrapod-bearing sites that produce taxa thought to possess an herbivorous diet. Table 4. Virgilian locality data, listing those tetrapod-bearing sites that produce taxa thought to possess an herbivorous diet. Table 5. Wolfcampian locality data, listing those tetrapod-bearing sites that produce taxa thought to possess an herbivorous diet. Table 6. Leonardian locality data, listing those tetrapod-bearing sites that produce taxa thought to possess an herbivorous diet. viii List of Figures Fig. 1. MCZ 9436, right maxilla and holotype of Ambedus pusillus in lateral (A), occlusal (B), and medial (C) views. Numbers indicate tooth positions. From Kissel and Reisz (2004a). Scale = 1 cm. Fig. 2. Fifth posteriormost maxillary tooth of MCZ 9436 (A) and fifth posteriormost dentary tooth of MCZ 9440 (B) in posterior view. From Kissel and Reisz (2004a). Scale = 1 cm. Fig. 3. MCZ 9438, complete left dentary of Ambedus pusillus in lateral view. Numbers indicate tooth positions. From Kissel and Reisz (2004a). Scale = 1 cm. Fig. 4. MCZ 9439, right dentary of Ambedus pusillus in lateral (A) and medial (B) views. Numbers indicate tooth positions. From Kissel and Reisz (2004a). Scale = 1 cm. Fig. 5. MCZ 9440, left dentary of Ambedus pusillus in lateral (A) and occlusal (B) views. Numbers indicate tooth positions. From Kissel and Reisz (2004a). Scale = 1 cm. Fig. 6. OMNH 56871, articulated left frontal and postfrontal in dorsal view (A), OMNH 56872a (B) and OMNH 56873 (C), premaxillary incisor teeth in lingual and lateral views, and OMNH 56875 (D) and OMNH 56872b (E) in probable posterior, anterior, and lingual views. From Reisz and Sutherland (2001). Scale = 1 cm. Fig. 7. Articulated series of four presacral and two sacral vertebrae of Phanerosaurus naumanni, the sole specimen assigned to the form, in left lateral (top) and dorsal (bottom) views. From Meyer (1860; no scale provided). Fig. 8. Partial left lower jaw in medial view (A), partial right dentary in medial view (B), right maxilla in lateral view (C), and vertebrae (D) of Stephanospondylus pugnax; MNG 14000, a partial skull of Orobates pabsti in lateral view (E). B from Stappenbeck (1905; no scale provided); C-D from Geinitz and Deichmüller (1882; no scale provided); scale for E = 1 cm. Fig. 9. Hypothesis demonstrating the monophyly of Diadectomorpha and Diadectidae. Autapomorphies of the clades and terminal taxa are as follows, with all characters listed possessing an unambiguous history: Node A (Cotylosauria): 5(1), 33(1), 35(1); Amniota: 1(1), ix 11(1), 24(1); Node B (Diadectomorpha): 2(1), 4(1), 18(1), 25(1), 26(1), 32(1), 36(1), 37(1); Node C: 3(1), 8(1); Node D (Diadectidae): 27(1), 30(1), 31(1); Node E: 22(1), 26(2), 28(1); Node F: 12(1); 20(1); Node G: 19(1), 22(2), 30(2), 31(2). Bootstrap and decay index values determined in Paup 4.0b10 are: Node B: 97, 4; Node C: 56, 1; Node D: 96, 3; Node E: 95, 3; Node F: 95, 3; Node G: 92, 3. Fig. 10. CM 1938, holotype and only known specimen (a partial left maxilla) of Desmatodon hollandi in lateral (A), medial (B), and occlusal (C) views. Scale = 1 cm. Fig. 11. Skull (palatal view; A) and right lower jaw of Diadectes lentus in occlusal (B), medial (C), and lateral (D) views. Scale = 1 cm. Ang, angular; s.ang, surangular; art, articular; p.art, prearticular; d, dentary; spl, splenial; ant fenestra, anterior fenestra; med fenestra, medial fenestra; post fenestra, posterior fenestra; numbers indicate tooth positions. A and B from Case and Williston (1912); C-E from Welles (1941). Fig. 12. YPM 817, partial skull of Diadectes carinatus in palatal view. From Case and Williston (1912). Scale = 1 cm. Fig. 13. Skull reconstruction of Limnoscelis paludis is dorsal (A), ventral (B), lateral (C), and occipital (D) views, as well as the lateral (E) and medial (F) views of the lower jaw, based on holotype YPM 811. Scale = 2 cm. a, angular; ac, anterior coronoid; ar, articular; bo, basioccipital; d, dentary; ec, ectopterygoid; f, frontal; j, jugal; l, lacrimal; m, maxilla; n, nasal; op, opisthotic; p, parietal; pc, posterior coronoid; pf, postfrontal; pl, palatine; pm, premaxilla; po, postorbital; pp, postparietal; pra, prearticular; prf, prefrontal; ps, parasphenoid; pt, pterygoid; q, quadrate; qj, quadratojugal; s, septomaxilla; sa, surangular; so, supraoccipital; sp, splenial; sq, squamosal; st, supratemporal; t, tabular; v, vomer.