Anatomia, Taxonomia, Ontogenia E Filogenia De Mosassaurianos Basais (Squamata, Mosasauria) E Suas Implicações Para a Evolução De Anguimorpha
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Bruno Gonçalves Augusta Anatomia, taxonomia, ontogenia e filogenia de mosassaurianos basais (Squamata, Mosasauria) e suas implicações para a evolução de Anguimorpha Anatomy, taxonomy, ontogeny and phylogeny of basal mosasaurians (Squamata, Mosasauria) and their implications to the evolution of Anguimorpha Vol. 2 – Figures São Paulo 2019 242 Chapter 1 - A new and exquisitely preserved fossil marine lizard with embryos Figure 1.1 – Coniasaurus sp. nov. preserved remains. Adult remains (above) are: A) Parietal; B) Cervical vertebra; C) Dorsal vertebra; D) Sacral vertebrae; E) Pygal vertebra; F) Dentary; G) Humerus; H) Proximal end of femur; I) Distal end of femur; J) Caudal vertebra. Embryonic remains (below) are: K) Premaxilla; L) Frontal; M) Cervical vertebra; N) Pygal vertebra; O) Dentary; P) Dorsal vertebra; Q) Caudal vertebra. Scale bars are 1mm for bones and 50mm for reconstructions. Skeletal reconstructions credit: Felipe Alves Elias. 243 Figure 1.2 – Paleogeography of Texas and biogeographic distribution of mosasaurians during the early Late Cretaceous (Cenomanian-Turonian). A) Cenomanian and Turonian dolichosaur and mosasaur localities. Southern North Sea Basin: 1: England and France (Anglo-Paris Basin; Rage, 1989; Caldwell, 1999; Caldwell & Cooper, 1999; 2: NW Germany (Diedrich, 1997, 1999). Eastern Mediterranean: 3: Slovenia (Carroll & DeBraga, 1992); 4: Hvar and Lesina, Croatia (Carroll & De Braga, 1992); 5: Hajula, Hakel and Al Nammoura, Lebanon (Dal Sasso & Renesto, 1999; Dalla Vecchia & Venturini, 1999); 6: ‘Ein Yabrud (Polcyn et al., 1999); 7: Besokty II, Kazakhstan (Averianov, 2001); 8: Goulmima, Morocco (Bardet et al., 2003); 9: Iembi, 244 Angola (Antunes, 1964); 10: Yagua, Columbia (Páramo, 1994). USA: 11: Big Bend, Texas (Bell & VonLoh, 1998); 12: Dallas area (Jacobs et al., 2005); 13: Kansas; 14: South Dakota and Wyoming (Bell & VonLoh, 1998; VonLoh & Bell, 1998). Buchy et al. (2005) report a plesiopedal mosasaur from Nuevo León, Mexico, near Big Bend. For a discussion of the age and correlation of these localities see Jacobs et al. (2005). Map after Hay et al. (1999). Figure after Jacobs et al 2005a Figure 1; B) Early Cenomanian, (Neogastroplites cornutus time ~97Ma); C) Late Cenomanian (Neocardioceras juddii time; ~94Ma); D) Late Turonian (Prionocyclus germari time ~89.85 Ma); B-D modified from Slattery et al., 2015. Figure 1.3 – Morphological comparison of Coniasaurus sp. nov. frontal bone. Columns I and II shows embryonic frontals, while column III depicts an adult one. Line A represent the bones in dorsal view and line B in ventral view. Scale bars: 1mm. 245 Figure 1.4 – Morphological comparison of Coniasaurus sp. nov. dentition. Dentary fragments are depicted in medial view, with embryonic specimens above and the adult below. Scale bars: 1mm. 246 Figure 1.5 – Phylogenetic relationships of the Dolichosauridae. Single tree recovered with 113 steps (CI: 0,72 and RI: 0,75). Numbers above nodes indicate values of Bremer supports, and numbers below it the Bootstrap values. Mosasauria and Mosasauroidea are indicated by square shapes. All non-mosasauroids mosasaurians in this tree are dolichosaurs. 247 Figure 1.6 – Aquatic adaptations of Coniasaurus sp. nov. A) Humerus showing a well- developed deltopectoral crest; B) Femur showing a specialized distal end; C) Caudal vertebra showing a very elongated haemal arch. Scale bars: 1cm. SUPPLEMENTARY FIGURES Figure S1 – Coniasaurus sp. nov. frontal fragment. A) Anterior view; B) Posterior view; C) Left side; D) Dorsal view; E) Ventral view; F) Right side. Scale bar: 2mm. 248 Figure S2 – Coniasaurus sp. nov. parietal. A) Anterior view; B) Posterior view; C) Left side; D) Dorsal view; E) Ventral view; F) Right side. Scale bar: 2mm. Figure S3 – Coniasaurus sp. nov. dentary. A) Lateral view; B) Medial view; C) Anterior view; D) Dorsal view; E) Ventral view; F) Posterior view. Scale bar: 2mm. 249 Figure S4 – Coniasaurus sp. nov. cervical vertebrae. Anteriormost element shown in: A) Anterior view; B) Posterior view; C) Left side; D) Dorsal view; E) Ventral view; F) Right side. Middle element shown in: G) Anterior view; H) Posterior view; I) Left side; 250 J) Dorsal view; K) Ventral view; L) Right side. Posteriormost element shown in: M) Anterior view; N) Posterior view; O) Left side; P) Dorsal view; Q) Ventral view; R) Right side. Scale bar: 2mm. Figure S5 – Coniasaurus sp. nov. dorsal vertebra. A) Anterior view; B) Posterior view; C) Left side; D) Dorsal view; E) Ventral view; F) Right side. Scale bar: 2mm. Figure S6 – Coniasaurus sp. nov. sacral vertebrae. A) Anterior view; B) Posterior view; C) Left side; D) CT horizontal section, evidencing the fusion of elements; E) Dorsal view; F) Ventral view; G) Right side; H) CT cross section, evidencing the fusion of elements. Scale bar: 2mm. 251 Figure S7 – Coniasaurus sp. nov. pygal vertebra. A) Anterior view; B) Posterior view; C) Left side; D) Dorsal view; E) Ventral view; F) Right side. Scale bar: 2mm. 252 Figure S8 – Coniasaurus sp. nov. caudal vertebra. A) Anterior view; B) Posterior view; C) Left side; D) Dorsal view; E) Ventral view; F) Right side. Scale bar: 2mm. 253 Figure S9 – Coniasaurus sp. nov. ribs. Cervical rib in: A) Dorsal view; B) Ventral view. Dorsal rib in: C) Dorsal view; D) Ventral view. Scale bar: 2mm. 254 Figure S10 – Coniasaurus sp. nov. humerus. A) Lateral view; B) Medial view; C) Proximal view; D) Anterior view; E) Posterior view; F) Distal view. Scale bar: 2mm. 255 Figure S11 – Coniasaurus sp. nov. femur. A) Lateral view; B) Medial view; C) Proximal view; D) Anterior view; E) Posterior view; F) Distal view. Scale bar: 2mm. 256 Figure S12 – Coniasaurus sp. nov. tibia. A) Lateral view; B) Medial view; C) Proximal view; D) Anterior view; E) Posterior view; F) Distal view. Scale bar: 2mm. 257 Figure S13 – Coniasaurus sp. nov. fibula. A) Lateral view; B) Medial view; C) Proximal view; D) Anterior view; E) Posterior view; F) Distal view. Scale bar: 2mm. 258 Figure S14 – Coniasaurus sp. nov. metatarsal element. A) Lateral view; B) Medial view; C) Proximal view; D) Anterior view; E) Posterior view; F) Distal view. Scale bar: 2mm. 259 Figure S15 – Coniasaurus sp. nov. phalanx. A) Lateral view; B) Medial view; C) Proximal view; D) Anterior view; E) Posterior view; F) Distal view. Scale bar: 2mm. 260 Figure S16 – Coniasaurus sp. nov. ungual. A) Lateral view; B) Medial view; C) Proximal view; D) Anterior view; E) Posterior view; F) Distal view. Scale bar: 2mm. 261 Figure S17 – Character 1 of the present analysis. Dentition, "coniasaur-like" heterodonty: (0) absent (Opetiosaurus buccichi); (1) present, with posteriormost teeth being significantly distinct in shape and proportions from the anteriormost teeth and bearing an anterodorsally oriented carena (Coniasaurus sp. nov.). Figure S18 – Character 2 of the present analysis. Dentition, presence of a conspicuous groove on the anterolateral surface of the crown: (0) absent (Opetiosaurus buccichi); (1) present (Coniasaurus sp. nov.). 262 Figure S19 – Character 16 of the present analysis. Frontal, proportional size: (0) short, frontal nearly as long as wide (Lanthanotus borneensis); (1) elongated, frontal much longer than wide (Opetiosaurus buccichi). State 0 image: modified from McDowell and Bogert (1954). State 1 image: modified from Dutchak and Caldwell (2009). Figure S20 – Character 18 of the present analysis. Parietal, width of dorsal aspect of parietal table: (0) very large anteriorly and posteriorly (Lanthanotus borneensis); (1) gradually tapers posteriorly (Coniasaurus sp. nov.); (2) very thin posteriorly (Pontosaurus lesinensi). State 0 image: modified from McDowell and Bogert (1954). 263 Figure S21 – Character 44 of the present analysis. Dorsal vertebrae, shape: (0) “V” shaped, with lateral sides converging towards condyle (Pontosaurus lesinensis); (1) “T” shaped, with abrupt expansion anteriorly (Coniasaurus sp. nov.); (2) “Box” shaped, with lateral sides running nearly parallel (Mosasaurus hoffmanni). State 2 image: modified from Street and Caldwell (2016). Figure S22 – Character 45 of the present analysis. Dorsal vertebrae, presence of a lateral “carving” in the lateral aspect of the vertebrae, between the centrum and the neural arch: (0) absent (Coniasaurus crassidens); (1) present (Coniasaurus sp. nov.). 264 Figure S23 – Character 46 of the present analysis. Dorsal vertebrae, width of posterior base of the neural spine: (0) equal or subequal to the anterior base (Coniasaurus crassidens); (1) posterior base larger than anterior one (Coniasaurus sp. nov.). Figure S24 – Character 48 of the present analysis. Sacral vertebrae, fusion: (0) absent (Coniasaurus longicollis); (1) present (Coniasaurus sp. nov.). Figure S25 – Character 49 of the present analysis. Caudal vertebrae, deepness: (0) regular (Varanus salvator); (1) very deep, forming a strong sculling organ (Opetiosaurus buccichi). 265 Figure S26 – Characters 58, 60 and 61 of the present analysis. 58. Humerus, deltopectoral crest proportional size: (0) larger length of the crest measuring less than 50% of the humerus shaft lenght; (1) larger length of the crest measuring 50% or more of the humerus shaft length. 60. Humerus, shaft twisted: (0) absent; (1) present. 61. Humerus, distinct tubercle in the mid-point of the shaft: (0) absent; (1) present. Image representing state 0: Acteosaurus tommasinii. Image representing state 1: Coniasaurus sp. nov. 266 Chapter 2 - New remains of North American coniasaurs (Squamata: Mosasauria) from the Upper Cretaceous of Texas and a revision of the genus Coniasaurus