Anatomy, taxonomy, ontogeny and phylogeny of basal mosasaurians (Squamata, Mosasauria) and their implications to the evolution of Anguimorpha

Texto

(1)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.

(2) 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..

(3) 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,.

(4) 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..

(5) 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..

(6) 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..

(7) 247. Figure 1.6 – Aquatic adaptations of Coniasaurus sp. nov. A) Humerus showing a welldeveloped 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..

(8) 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..

(9) 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;.

(10) 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..

(11) 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..

(12) 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..

(13) 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..

(14) 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..

(15) 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..

(16) 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..

(17) 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..

(18) 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..

(19) 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..

(20) 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..

(21) 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.)..

(22) 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)..

(23) 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.)..

(24) 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)..

(25) 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..

(26) 266. Chapter 2 - New remains of North American coniasaurs (Squamata: Mosasauria) from the Upper Cretaceous of Texas and a revision of the genus Coniasaurus. Figure 2.1 – Location map and Eagle Ford outcrop area. After Kennedy, 1988..

(27) 267. Figure 2.2 – Generalized sections of the Eagle Ford Shale in Dallas area and a summary of Coniasaurus producing localities. Data from: Adkins and Lozo, 1951; McNulty, 1954; Brown and Pierce 1962; McNulty, 1966; Smith. 1981; Kennedy, 1988; Kennedy and Cobban,1990; Hancock and Walaszczyk, 2004..

(28) 268. Figure 2.3 – Generalized sections of the Eagle Ford Shale in the Temple-Austin area and a summary of Coniasaurus producing localities. Data from: Adkins and Lozo, 1951, Kennedy and Cobban, 1990..

(29) 269. Figure 2.4 – Coniasaur (SMU-72396) cervical vertebrae. A more anterior element exposed in: A) Anterior view; B) Posterior view; C) Left side; D) Dorsal view; E) Ventral view; F) Right side; and a more posterior element exposed in: G) Anterior view; H) Posterior view; I) Left side; J) Dorsal view; K) Ventral view; L) Right side. Scale bar: 1mm..

(30) 270. Figure 2.5 – Coniasaur (SMU-72396) dorsal vertebrae. Single isolated element exposed in: A) Anterior view; B) Posterior view; C) Left side; D) Right side; E) Dorsal view; F) Ventral view. Articulated elements exposed in: G) Anterior view; H) Posterior view; I) Dorsal view; J) Left side; K) Ventral view; L) Right side. Scale bar: 2mm..

(31) 271. Figure 2.6 – Coniasaur (SMU-72396) sacral vertebrae and ribs. A) Anterior view; B) Posterior view; C) Left side; D) Dorsal view; E) Ventral view; F) Right side; G) Proximal fragment of a sacral rib; H) Distal fragment of a sacral rib. Scale bar: 2mm.. Figure 2.7 – Coniasaur (SMU-72396) pygal vertebra. A) Anterior view; B) Posterior view; C) Left side; D) Right side; E) Ventral view; F) Right side. Scale bar: 2mm..

(32) 272. Figure 2.8 – Coniasaur (SMU-72396) caudal vertebra. A) Anterior view; B) Posterior view; C) Left side; D) Right side; E) Ventral view; F) Right side. Scale bar: 2mm.. Figure 2.9 – Coniasaur (SMU-72396) ilium. A) Lateral view; B) Medial view. Scale bar: 2mm..

(33) 273. Figure 2.10 – Coniasaur maxillary fragments from the DFW Airport locality. SMU72384 in: A) Lateral view; B) Medial view; C) Dorsal view; D) Ventral view. SMU72391 in: E) Lateral view; F) Medial view; G) Dorsal view; H) Ventral view. Scale bar: 2mm.. Figure 2.11 – Coniasaur (SMU-72389) frontal fragment from the DFW Airport locality. A) Anterior view; B) Posterior view; C) Left side; D) Dorsal view; E) Ventral view; F) Right side. Scale bar: 2mm..

(34) 274. Figure 2.12 – Coniasaur dentary fragments from the DFW Airport locality. SMU72385 in: A) Lateral view; B) Medial view. SMU-72386 in: C) Lateral view; D) Medial view. Scale bar: 2mm.. Figure 2.13 – Coniasaur (SMU-72382) surangular. A) Anterior view; B) Posterior view; C) Lateral view; D) Dorsal view; E) Ventral view; F) Medial view. Scale bar: 2mm..

(35) 275. Figure 2.14 – Coniasaur (SMU-76276) dentary from the Lowes Farm location. A) Lateral view; B) Medial view; C) Dorsal view; D) Ventral view. Scale bar: 4mm..

(36) 276. Figure 2.15 – Coniasaur (SMU-72416) humerus from the New Locality. A) Anterior view; B) Posterior view; C) Proximal view; D) Lateral view; E) Medial view; F) Distal view. Scale bar: 2mm..

(37) 277. Figure 2.16 – Coniasaur (TMM 40305-1) maxilla. A) Lateral view; B) Medial view; C) Dorsal view; D) Ventral view. Scale bar: 4mm.. Figure 2.17 – Coniasaur maxillary fragments from the DFW Airport locality cited in Bell et al. (1982). SMU-69020 first fragment in A) Lateral view; B) Medial view; and second fragment in: C) Lateral view; D) Medial view. SMU-69021 in: E) Lateral view; F) Medial view. SMU-69023 in: G) Lateral view; H) Medial view. Scale bar: 2mm..

(38) 278. Figure 2.18 – Coniasaur (SMU-69028) pterygoid from the DFW Airport locality cited in Bell et al. (1982). A) Anterior view; B) Posterior view; C) Lateral view; D) Dorsal view; E) Ventral view; F) Medial view. Scale bar: 2mm.. Figure 2.19 – Coniasaur dentary fragments from the DFW Airport locality cited in Bell et al. (1982). SMU-69019 in: A) Lateral view; B) Medial view. SMU-69020 in: C) Lateral view; D) Medial view; SMU-69027 in: E) Lateral view; F) Medial view. Scale bar: 2mm..

(39) 279. Figure 2.20 – Adriosaurus (NHMUK-R2867) skull, with special reference to frontal anatomy. A) Line drawing of the specimen provided by Lee and Caldwell (2000); B) Photograph of the same specimen; C) Same photograph, but with frontal highlighted by black lines..

(40) 280. Chapter 3 - Patterns of tooth attachment and implantation in Coniasaurus (Squamata, Mosasauria) based on highresolution CT data. Figure 3.1 – General patterns of tooth attachment and implantation in Coniasaurus. Tooth-bearing bones are depicted in medial view. A) Left dentary, DMNH-1601; B) Left dentary, SMU-72398; C) Left maxilla of an embryonic specimen, DMNH-1601; D) Left maxilla, SMU-72390. Scale bar = 2mm..

(41) 281. Figure 3.2 – Tooth attachment tissues in Coniasaurus. A) Longitudinal section of a left maxilla, SMU-72390; B) Detail of the same section near the tooth base; C) Longitudinal section of a left dentary, SMU-72398; D) Sagittal section of a left maxilla, SMU-72390. Abbreviations: AB = Alveolar Bone; CEM = Cementum..

(42) 282. Figure 3.3 – Dentin infoldings in Coniasaurus. Arrow points to the tooth element from which cross sections were taken, and lines indicate the level of cross sections. A) Left dentary, SMU-72398; B) Upper cross section of the indicated element; C) Lower cross section of the indicated element..

(43) 283. Chapter 4 - Ontogenetic trends in Coniasaurus (Squamata, Mosasauria) and its implications to the paleobiology and evolution of basal mosasaurians. Figure 4.1 – Coniasaurus sp. nov. (DMNH-1601) embryonic premaxilla. A) Dorsal view; B) Ventral view; C) Left side; D) Anterior view; E) Right side; F) Pontosaurus lesinensis premaxilla (note the differences regarding the narial bar thickness). Scale bar = 1mm..

(44) 284. Figure 4.2 – Frontal ontogenetic trends in Coniasaurus. Columns indicate specimens as follows: I) Coniasaurus sp. nov. embryonic frontal (DMNH-1601); II) Coniasaurus sp. nov. embryonic frontal (DMNH-1601); III) Coniasaurus sp. nov. adult frontal (DMNH-1601); IV) C. gracilodens frontal (NHMUK-R44141). Lines indicate views as follows: A) Dorsal; B) Ventral; C) Anterior; D) Posterior; E) Left side; F) Right side. Scale bars = 1mm..

(45) 285. Figure 4.3 – Coniasaurus sp. nov. (DMNH-1601) embryonic quadrate. A) Medial view; B) Lateral view; C) Dorsal view; D) Anterior view; E) Posterior view; F) Ventral view. Scale bar = 1mm..

(46) 286. Figure 4.4 – Maxilla ontogenetic trends in Coniasaurus. Columns indicate specimens as follows: I) Coniasaurus sp. nov. embryonic maxilla (DMNH-1601); II) Coniasaurus spp. juvenile maxilla (SMU-72391); III) Coniasaurus spp. adult maxilla (SMU-72390). Lines indicate views as follows: A) Medial; B) Lateral; C) Dorsal; D) Ventral. Scale bars = 1mm.. Figure 4.5 – Dentary ontogenetic trends in Coniasaurus. Columns indicate specimens as follows: I) Coniasaurus sp. nov. embryonic dentary fragments (DMNH-1601); II).

(47) 287 Coniasaurus spp. juvenile dentary (SMU-72398); III) Coniasaurus sp. nov. adult dentary (DMNH-1601). Lines indicate views as follows: A) Medial; B) Lateral; C) Dorsal; D) Ventral. Scale bars = 1mm.. Figure 4.6 – Dentition ontogenetic trends in Coniasaurus. Teeth are always shown in groups of three elements, from the left to the right: a more anterior, mid- and a more posterior element. Columns indicate specimens as follows: I) Coniasaurus sp. nov. embryonic teeth (DMNH-1601); II) Coniasaurus spp. juvenile teeth (SMU-72391 and SMU-72398); III) Coniasaurus sp. nov. adult teeth (DMNH-1601). Lines indicate views as follows: A) Medial; B) Lateral; C) Occlusal. Scale bars = 1mm..

(48) 288. Figure 4.7 – Cervical vertebrae ontogenetic trends in Coniasaurus. Columns indicate specimens as follows: I) Coniasaurus sp. nov. embryonic cervical (DMNH-1601); II) Coniasaurus spp. juvenile cervical (SMU-72396); III) Coniasaurus sp. nov. adult cervical (DMNH-1601); IV) Coniasaurus spp. adult cervical (SMU-69026). Lines indicate views as follows: A) Dorsal; B) Ventral; C) Left side; D) Anterior; E) Posterior. Scale bars = 1mm..

(49) 289. Figure 4.8 – Dorsal vertebrae ontogenetic trends in Coniasaurus. Columns indicate specimens as follows: I) Coniasaurus sp. nov. embryonic cervical (DMNH-1601); II) Coniasaurus spp. juvenile cervical (SMU-72396); III) Coniasaurus sp. nov. adult cervical (DMNH-1601); IV) Coniasaurus spp. adult cervical (SMU-69022). Lines indicate views as follows: A) Dorsal; B) Ventral; C) Left side; D) Anterior; E) Posterior. Scale bars = 1mm..

(50) 290. Figure 4.9 – Sacral vertebrae ontogenetic trends in Coniasaurus. Columns indicate specimens as follows: I) Coniasaurus spp. juvenile sacrals (SMU-72396); II) Coniasaurus sp. nov. adult sacrals (DMNH-1601. Lines indicate views as follows: A) Dorsal; B) Ventral; C) Left side; D) Anterior; E) Posterior. Scale bars = 1mm..

(51) 291. Figure 4.10 – Pygal vertebrae ontogenetic trends in Coniasaurus. Columns indicate specimens as follows: I) Coniasaurus sp. nov. embryonic pygal (DMNH-1601); II) Coniasaurus spp. juvenile pygal (SMU-72396); III) Coniasaurus sp. nov. adult pygal (DMNH-1601). Lines indicate views as follows: A) Dorsal; B) Ventral; C) Left side; D) Anterior; E) Posterior. Scale bars = 1mm..

(52) 292. Figure 4.11 – Caudal vertebrae ontogenetic trends in Coniasaurus. Columns indicate specimens as follows: I) Coniasaurus sp. nov. embryonic caudal (DMNH-1601); II) Coniasaurus spp. juvenile caudal (SMU-72396); III) Coniasaurus sp. nov. adult caudal (DMNH-1601); IV) Coniasaurus spp. adult caudal (SMU-69025). Lines indicate views as follows: A) Dorsal; B) Ventral; C) Left side; D) Anterior; E) Posterior. Scale bars = 1mm..

(53) 293. Figure 4.12 – Humerus ontogenetic trends in Coniasaurus. Columns indicate specimens as follows: I) Coniasaurus sp. nov. embryonic humerus, still attached to an adult vertebra (DMNH-1601); II) Coniasaurus spp. juvenile humerus (SMU-72416); III) Coniasaurus sp. nov. adult humerus (DMNH-1601). Lines indicate views as follows: A) Lateral; B) Medial; C) Anterior; D) Posterior; E) Proximal; F) Distal. Scale bars = 1mm..

(54) 294. Figure 4.13 – Isolated elements used to infer the number of embryonic individuals. Specimens belong to DMNH-1601, the holotype of Coniasaurus sp. nov. A and B are left dentaries, while C, D and E represent pygal vertebrae. Scale bars = 1mm..

(55) 295. Chapter 5 - Evolution of the postcranial skeleton in basal mosasaurians (Squamata, Mosasauria). Figure 5.1 – Cervical vertebrae of basal mosasaurians. A) Atlas and axis of Eidolosaurus trauthi outlined, ventral view, based on a cast of the holotype (NMWunnumbered); B) 3D reconstruction of a Coniasaurus sp. nov. (DMNH-1601) cervical element, lateral view, arrow pointing to the fused hypapophyses-intercentrum complex; C) Detail of cervical vertebrae in Pontosaurus lesinensis (GBA-1873), ventral view, arrow pointing to the unfused hypapophyses-intercentrum complex. Scale bar = 5mm..

(56) 296. Figure 5.2 – Dorsal vertebrae of basal mosasaurians. A) Coniasaurus sp. nov. (DMNH-1601), dorsal view; B) Adriosaurus suessi (NHMUK PR 2867), dorsal view; C) Pontosaurus lesinensis (GBA-1873), ventral view; D) Komensaurus carrolli (MCSNT11430-2), ventral view. Scale bar = 5mm..

(57) 297. Figure 5.3 – Sacral vertebrae of basal mosasaurians and Heloderma. A) Coniasaurus sp. nov. (DMNH-1601), dorsal view; B) Coniasaurus longicollis (BMB 0085687), ventral view; C) Heloderma (SMU unnumbered), ventral view. Scale bar = 5mm..

(58) 298. Figure 5.4 – Detail of Adriosaurus suessi manus (NHMUK PR 2867). Scale bar = 5mm.. Figure 5.5 – Humerus of basal mosasaurians. A) Adriosaurus microbrachis right humerus (MCSNT 7792); B) Adriosaurus microbrachis left humerus (same specimen); C) Coniasaurus sp. nov. (DMNH-1601). Scale bar = 5mm..

(59) 299. Figure 5.6 – Pubis of basal mosasaurians and marine fossil snakes. A) Adriosaurus sp. (MCSNT-11426); B) Komensaurus carrolli (MCSNT-11430-2); C) Reconstruction of Pachyrhachis problematicus pelvis, modified from Lee and Caldwell (1998); D) 3D reconstruction of Eupodophis descouensi, modified from Houssaye et al. (2011). Scale bar = 5mm..

(60) 300. Figure 5.7 – Ilium interpretations of Adriosaurus suessi (NHMUK PR 2867). A) Interpretative drawing of the pelvic region from Lee and Caldwell (2000); B) Photograph of the specimen pelvic region; C) Same photograph, but with ilium outlined for facilitating visualization. Scale bar = 5mm..

(61) 301. Figure 5.8 – Detail of Coniasaurus longicollis femur (BMB 0085687). Scale bar = 5mm.. Figure 5.9 – Tibia of basal mosasaurians, with white arrows pointing to the notch present on the distal tibial ends. A) Coniasaurus sp. nov. (DMNH-1601); B) Komensaurus carrolli (MCSNT-11430-2). Scale bar = 5mm..

(62) 302. Chapter 6 - A phylogenetic analysis of Mosasauria (Squamata) supports the hypothesis of varanoid affinities for the clade. Figure 6.1 (Part 1) – Phylogenetic relationships of squamates. Strict consensus of 100.000 MPTs. Continued on next page..

(63) 303. Figure 6.1 (Part 2) – Phylogenetic relationships of squamates. Strict consensus of 100.000 MPTs. Continued on next page..

(64) 304. Figure 6.1 (Part 3) – Phylogenetic relationships of squamates. Strict consensus of 100.000 MPTs. Mosasauria is highlighted by the square shape..

(65) 305. Figure 6.2 – Phylogenetic relationships of squamates, showing here only anguimorphs, after removing Judeasaurus and Pannoniasaurus. Strict consensus of 100.000 MPTs (CI: 0,185; RI: 0,793). Numbers above nodes indicate values of Bremer supports, and numbers below it the Bootstrap values. Mosasauria, Mosasauroidea and Dolichosauridae are indicated by square shapes..

(66) 306. Figure 6.3 – Phylogenetic relationships of squamates, showing here only anguimorphs, after removing Judeasaurus and Pannoniasaurus and turning off character ordering. Strict consensus of 100.000 MPTs and 4866 steps. Mosasauria is highlighted by the square shape..

(67) 307. Figure 6.4 – Phylogenetic relationships of squamates, showing here only anguimorphs, after taxonomic test 1: including only dolichosaurs. Strict consensus of 13104 MPTs and 5099 steps. Mosasauria is highlighted by the square shape..

(68) 308. Figure 6.5 – Phylogenetic relationships of squamates, showing here only anguimorphs, after taxonomic test 2: including only basal mosasauroids. Strict consensus of 15528 MPTs and 5080 steps. Mosasauria is highlighted by the square shape..

(69) 309. Figure 6.6 – Phylogenetic relationships of squamates, showing here only anguimorphs, after taxonomic test 3: including only mosasaurids. Strict consensus of 23040 MPTs and 5218 steps. Mosasauria is highlighted by the square shape..

(70) 310. Figure 6.7 – Phylogenetic relationships of squamates, showing here only anguimorphs, after taxonomic test 4: including only hydropedal mosasaurids. Strict consensus of 7680 MPTs and 5178 steps. Mosasauria is highlighted by the square shape..

(71) 311. Figure 6.8 – Phylogenetic relationships of squamates, showing here only anguimorphs, after taxonomic test 5: removing dolichosaurs. Strict consensus of 100.000 MPTs and 5242 steps. Mosasauria is highlighted by the square shape..

(72) 312. Figure 6.9 – Phylogenetic relationships of squamates, showing here only anguimorphs, after taxonomic test 6: removing basal mosasauroids. Strict consensus of 40132 MPTs and 5271 steps. Mosasauria is highlighted by the square shape..

(73) 313. Figure 6.10 – Phylogenetic relationships of squamates, showing here only anguimorphs, after taxonomic test 7: removing mosasaurids. Strict consensus of 36512 MPTs and 5130 steps. Mosasauria is highlighted by the square shape..

(74) 314. Figure 6.11 – Phylogenetic relationships of squamates, showing here only anguimorphs, after taxonomic test 8: removing hydropedal mosasaurids. Strict consensus of 100.000 MPTs and 5173 steps. Mosasauria is highlighted by the square shape..

(75) 315. Figure S1 – Character 617 of the present analysis. Humerus, deltopectoral crest shape: (0) rounded; (1) squared.. Figure S2 – Character 619 of the present analysis. Radius and ulna: (0) Parallel or subparallel in relation to each other (Shinisaurus crocodilurus); (1) divergent elements (Clidastes). State 0 image: modified from Conrad (2006). State 1 image: modified from Russell (1967)..

(76)