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Figure 9 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 9. Photographs and line drawings of the posterior cervical vertebra MCF-PVPH-261/01 of Ligabuesaurus leanzai in anterior (A), posterior (B) and left lateral (C) views. Abbreviations: aas, anterior articular surface; apf, anterior pneumatic fossa; cprl, centroprezygapophyseal lamina; di, diapophysis; pa, parapophysis; pas, posterior articular surface; ppf, posterior pneumatic fossa; prz, prezygapophysis; r?, fragment of rib. Scale bar: 10 cm.

Figure 11 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 11. Photographs and line drawings of the posterior cervical vertebra MCF-PVPH-228/01 and MCF-PVPH-228/02 of Ligabuesaurus leanzai in anterior (A) view. Abbreviations: aas, anterior articular surface; cpol, centropostzygapophyseal lamina; di, diapophysis; poz, postzygapophysis; prdl, prezygodiapohyseal lamina; prz, prezygapophysis; sprl, spinoprezygapophyseal lamina; tprl, intraprezygapophyseal lamina. Scale bar: 10 cm.

Figure 10 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 10. Photographs and line drawings of the posterior cervical vertebra MCF-PVPH-261/02 of Ligabuesaurus leanzai in anterior (A), posterior (B) and right lateral (C) views. Abbreviations: apf, anterior pneumatic fossa; cprl, centroprezygapophyseal lamina; di, diapophysis; nc, neural canal; pa, parapophysis; ppf, posterior pneumatic fossa; prz, prezygapophysis. Scale bar: 10 cm.

Figure 8 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 8. Photographs and line drawings of the middle cervical vertebra MCF-PVPH-261/16 of Ligabuesaurus leanzai in dorsal (A), ventral (B) and left lateral (C) views. Abbreviations: acdl, anterior centrodiapophyseal lamina; apo, accessory pneumatic opening; cpol, centropostzygapophyseal lamina; cprl, centroprezygapophyseal lamina; di, diapophysis; nc, neural canal; pa, parapophysis; pcdl, posterior centrodiapophyseal lamina; pl, pleurocoel; prdl, prezygodiapophyseal lamina; prz, prezygapophysis; pvr, posteroventral ridges; spdl, spinodiapophyseal lamina; sprl, spinoprezygapophyseal lamina. Scale bar: 10 cm.

Figure 7. A in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 7. A, detail of the microwear pattern on the apical wear facet of the maxillary tooth MCF-PVPH-744 of Ligabuesaurus leanzai. Most of the pits and coarse grooves are evident with a binocular microscope (α) on the dentine exposed surface, whereas the scanning electron photomicrographs (β, γ, δ and ε) show abundant fine scars in both the enamel and dentine surfaces. Abbreviations: g, coarse groove; p, pit; s, fine scar.

Figure 5 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 5. New autapomorphic feature and revised diagnosis of Ligabuesaurus leanzai. A, Photograph and line drawings of the left coracoid MCF-PVPH-261/05 in lateral view. The quadrangular ventral half of the coracoid in lateral view is here proposed as a uniquely derived feature of Ligabuesaurus leanzai not present in any other sauropods. B–K, comparative outlines of sauropod coracoids: B, Neuquensaurus MLP-Ly-14; C, Saltasaurus PVL-4017-100; D, Quetecsaurus UNCUYO-LD-300.15; E, Opisthocoelicaudia ZPAL-MgD-I/48; F, Dreadnoughtus MPM-PV-1156; G, Tapuiasaurus MZSP-PV-807; H, Uberabatitan CPP-1109-UrHo; I, Suuwassea ANS 21122; J, Patagotitan MPEF-PV-3400/24; K, Zby ML 368. L–N, line drawings of the posterior cervical vertebra MCF-PVPH-233/02 (L) and anterior dorsal vertebrae MCF-PVPH-233/03 (M) and MCF-PVPH-908 (N), showing the development of the prespinal lamina on the anterior surface of the neural spine (green triangle). B–K modified from González Riga et al. (2019). Not to scale. Scale bar: 10 cm in L–N.

Figure 4 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 4. The somphospondylan sauropod Ligabuesaurus leanzai from Cerro de los Leones (Neuquén Province, Patagonia, Argentina). A, skeletal reconstruction based on the holotype (MCF-PVPH-233) and referred material (MCF-PVPH-261) from quarry no. 4, plus the newly referred specimens from quarry no. 3 (MCF-PVPH-228, MCF-PVPH-744 and MCF-PVPH-908). B, life restoration of Ligabuesaurus leanzai as a non-titanosaurian somphospondylan by J. L. Blanco.

Figure 3 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 3. Original picture of quarry no. 4 during the fieldwork of 2004 (A) and a quarry map with tentative arrangement of the holotype (in red) and referred material (in blue) of Ligabuesaurus leanzai (B). Abbreviations: ant, anterior; s/n, without collection number; v, vertebra.

Figure 27 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 27. Time-calibrated phylogenetic tree, showing geographical distribution and chronostratigraphical range of Somphospondyli, based on the agreement subtree of the reduced strict consensus. Lognkosauria and Lithostrotia have been collapsed into a single lineage. Ligabuesaurus leanzai (black narrow), from the Early Cretaceous Lohan Cura Formation (Neuquén Basin), was recovered as a non-titanosaurian somphospondylan.

Figure 2. Lithological profile. A in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 2. Lithological profile. A schematic log of the the lower section of the Cullin Grande Member (Bajada del Agrio Group, Lohan Cura Formation, Lower Cretaceous, Albian) that outcrops at the Cerro de los Leones locality (modified from Martinelli et al., 2007). Abbreviations: CS, crevasse channel; FF, floodplain fines; FL, fossiliferous level; LA, lateral accretion; LS, laminated sand sheets; LV, levee; SB, sandy bedforms. Architectural element codes follow Miall (1996).

Figure 1. Location map. A in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 1. Location map. A, satellite map of northern Patagonia (Argentina), showing the approximate limits of Neuquén Basin (white dashed line). B, the Cerro de los Leones locality is ~10 km to the south-west of Picún Leufú city, to the south of the Neuquén Province (white crossed pickaxes). C, satellite map of the main quarries opened on the southern flank of the Cerro de los Leones, where outcropping of the fluvial sediments of the Lohan Cura Formation (Albian) occurs.

Figure 15 in Anatomy and relationships of the bizarre Early Cretaceous pliosaurid Luskhan itilensis

Figure 15. Comparative cranial anatomy of raptorial marine amniotes with procumbent mesialmost teeth. All are high-resolution three-dimensional models. A, Luskhan itilensis (YKM 68344/1_262). B, PerƲushoƲisaurus campylodon (CAMSM B20671). C, Prognathodon solƲayi (IRSNB R33b). D, Ninoziphius platyrostris (MNHN SAS 941). E, Tasmacetus shepherdi (USNM 484878). F, Ziphius caƲirostris (NHMUK 1915.7.20.1). G, Ankylorhiza tiedemanni (CCNHM 103). H, Prosqualodon daƲidis (USNM 467596). Silhouettes: Ziphius by Chris Huh, from Phylopic (http://phylopic.org/ image/7c1d06fb-2d6e-454d-b57b-a859d5dbdb9f/); Squalodon by Craig Hylke, from Phylopic (http://phylopic.org/ image/95a73c63-e7c7-4e81-8e6d-592f647b07bc/).

Figure 3 in Anatomy and relationships of the bizarre Early Cretaceous pliosaurid Luskhan itilensis

Figure 3. Craniomandiobular osteology of the holotype of Luskhan itilensis (YKM 68344/1_262), based on a highresolution three-dimensional model. A, B, model (A) and interpretation (B) of the skull in left lateral view. C, D, model (C) and interpretation (D) of the skull in ventral view. Uncertain sutures are indicated by a dotted line.

Figure 2 in Anatomy and relationships of the bizarre Early Cretaceous pliosaurid Luskhan itilensis

Figure 2. Craniomandiobular osteology of the holotype of Luskhan itilensis (YKM 68344/1_262). A, B, photograph (A) and interpretation (B) of the skull in dorsal view. C–F, image and interpretation of four cross-sections (locations indicated in A). Uncertain sutures are indicated by a dotted line. The left postorbital bar and temporal arch are present but have not been photographed; they are present in Figure 3.

Figure 12 in Anatomy and relationships of the bizarre Early Cretaceous pliosaurid Luskhan itilensis

Figure 12. Pelvic girdle of the holotype of Luskhan itilensis (YKM 68344/1_262). A–D, left ilium in dorsal (A), medial (B), ventral (C) and lateral (D) views, along with a reconstruction of the possible extent of the dorsal blade (dotted line). E, F, right ilium in dorsal (E) and anterolateral (F) views.

Figure 9 in Anatomy and relationships of the bizarre Early Cretaceous pliosaurid Luskhan itilensis

Figure 9. Axial skeleton of the holotype of Luskhan itilensis (YKM 68344/1_262) based on high-resolution threedimensional models. A, B anterior cervical vertebrae in left lateral (A) and ventral (B) views. C–G, dorsal centra in dorsal (C), left lateral (D), anterior (E), right lateral (F) and ventral (G) views. H–M, right cervical rib in anterior (H), proximal (I), ventral (J), distal (K), posterior (L) and dorsal (M) views. N–R, cervical neural arch in right lateral (N), anterior (O), dorsal (P), left lateral (Q) and posterior (R) views. S–W, dorsal neural arch in anterior (S), ventral (T), left lateral (U), dorsal (V) and posterior (W) views. X–Z, caudal centra in dorsal (X), left lateral (Y) and ventral (Z) views. AA–AE, more posterior dorsal neural arch in anterior (AA), ventral (AB), left lateral (AC), dorsal (AD) and posterior (AE) views. AF–AH, dorsal rib in anterior (AF), proximal (AG) and posterior (AH) views.

Figure 6 in Anatomy and relationships of the bizarre Early Cretaceous pliosaurid Luskhan itilensis

Figure 6. Basicranium of the holotype of Luskhan itilensis (YKM 68344/1_262). Photographs and interpretations of the basicranium in posterodorsal (A, B), ventral (C, D) and left lateral (E, F) views.

Figure 11 in Anatomy and relationships of the bizarre Early Cretaceous pliosaurid Luskhan itilensis

Figure 11. Limbs of the holotype of Luskhan itilensis (YKM 68344/1_262). A, left forefin in dorsal view. B, C, right humerus in proximal (B) and distal (C) views. D, right forefin in dorsal view. E, F, right hind fin in ventral (E) and proximal (F) views. G, H, photographs and interpretation of the right hind fin in dorsal view.

Figure 8 in Anatomy and relationships of the bizarre Early Cretaceous pliosaurid Luskhan itilensis

Figure 8. Cervical region and centrum proportions of the holotype of Luskhan itilensis (YKM 68344/1_262). A, B, photographs and interpretation of the atlas–axis complex in left lateral view. C, D, photographs and interpretation of the cervical and pectoral regions, in dorsal view. E, evolution of the height-to-length ratio along the entire preserved vertebral column. Abbreviations: c, cervical; p, pectoral.

Figure 1. Luskhan itilensis, a in Anatomy and relationships of the bizarre Early Cretaceous pliosaurid Luskhan itilensis

Figure 1. Luskhan itilensis, a remarkably complete pliosaurid from the Lower Cretaceous of Russia. A, display of the holotype (YKM 68344/1_262) in the YKM. B, C, three-dimensional reconstruction of the skeleton of Luskhan itilensis in dorsal (B) and anterolateral (C) views. Please do not use this artistic reconstruction to describe anatomy or score phylogenetic characters.