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Fig. 2 in Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain

Fig. 2. Theropod dinosaurs teeth from upper Campanian–lower Maastrichtian, Laño. A, D. Theropoda indet. Morphotype 2. A. MCNA 14522. D. MCNA 1853. B, C.?Pyroraptor olympius Allain and Taquet, 2000. B. MCNA 14623. C. MCNA 14624. E–H. Theropoda indet. Morphotype 1. E. MCNA 1852. F. MCNA 14520. G. MCNA 14521. H. MCNA 2205. All lateral views. Scale bars 5 mm.

Fig. 5. A in Quadrupedal dinosaurs did not evolve fully pronated forearms: New evidence from the ulna

Fig. 5. A demonstration of the effects that changes in pronation of the distal radial epiphysis would have on manual orientation in a fully pronated tetrapod (the Virginia opossum Didelphis virginiana Kerr, 1792) that is using parasagittal forelimb kinematics with elbows inturned to the body wall. A. An unpronated forearm and manus, which if possible would point the manual digits posteriorly. B. A semi-pronated forearm and manus, which would point the manual digits laterally, analogous to many semi-pronated archosaurs, such as dinosaurs, that utilized quadrupedalism. C. A normal (for D. virginiana) fully pronated forearm and manus, which points the manual digits anteriorly. This demonstration illustrates why it was traditionally assumed that amniotes that evolve posteriorly directed elbows (i.e., archosaurs, therians and chameleons) would require an additional 90° of forearm pronation past the plesiomorphic 90°, because 180° of pronation is required to keep the wrist and finger joints aligned posteriorly during flexion, and therefore to continue operating in a parasagittal plane. Bonnan's (2003) radial hypothesis states that the radii of quadrupedal dinosaurs pronated to condition C, but the findings of this study show that the ulnae in these dinosaurs experienced an opposing supination that would have kept the planes of the wrist and finger joints wholly in condition in B, albeit with a tubular manus (e.g., Fig. 2E); see text for further discussion. The D. virginiana forelimb elements are from an adult specimen (FMNH 166984).

Fig. 11. A in Quadrupedal dinosaurs did not evolve fully pronated forearms: New evidence from the ulna

Fig. 11. A comparison of semi-pronated forearm views and distal antebrachial epiphyses of select pareiasaurs and synapsids. A. Pareiasaur, Embrithosaurus schwarzi Watson, 1914 (AMNH 2451), reversed. B. Pelycosaur, Dimetrodon loomisi Romer, 1937 (AMNH 21293), reversed, in flexor (B1) and distal (B2) views. C. Dinocephalian, Moschops capensis Broom, 1911 (AMNH 23930), reversed. D. Large dicynodont (AMNH 24096), reversed. E. Dinocephalian, Jonkeria haughtoni Broom, 1929 (AMNH 5577). F. Small dicynodont (uncataloged USNM), reversed. G. Dicynodont, Kannemeyeria simocephalus Weithofer, 1888 (AMNH 5591-93). Note that all specimens examined of the clades above possessed similar amounts of post-axial torsion of the distal ulnar diaphysis. Specimen in A oriented after Bradysaurus baini Seeley, 1892 ([FMNH] UC 1533 and UC 1525); specimens in C, F, and G after small dicynodonts Diictodon cf. grimbeeki (Broom, 1935) (USNM 412381 and USNM 452057). Not to scale.

Fig. 7 in Quadrupedal dinosaurs did not evolve fully pronated forearms: New evidence from the ulna

Fig. 7. Comparison of true standardized views of semi-pronated forearms in a representative sample of extant, nontherian tetrapods, in flexor (A1–F1) and distal (A2–F2) views. A. An allogatorid crocodilian, Alligator mississippiensis (Daudin, 1802) (FMNH 284695). B. A ratite bird, Struthio camelus Linnaeus, 1758 (FMNH 489294). C. A monitor lizard, Varanus komodoensis Ouwens, 1912 (FMNH 22197). D. A salamander, Ambystoma tigrinum (Green, 1825) FMNH 22010). E. A semi-aquatic turtle Apalone spinifera (Lesueur, 1827) (HDW NIU 1086), reversed. F. A more terrestrial turtle, Chrysemys picta (Schnei- der, 1783) (INHS 23894). Reversed specimens in this and following figures refer to elements from the right sides that have been digitally flipped. Note that in E the radius and ulna are fused in the morphology shown, and that the humerus may not be rotated far enough to the left. Note also that, in vivo, articular cartilage and, in some cases wrist bones (e.g., intermedium), may separate the distal radial and ulnar epiphyses beyond what is pictured here for specimens C and F. Not to scale.

Fig. 4. A in Quadrupedal dinosaurs did not evolve fully pronated forearms: New evidence from the ulna

Fig. 4. A demonstration of the effects that changes in pronation of the distal radial epiphysis would have on manual orientation in a semi-pronated tetrapod (Alligator mississippiensis [Daudin, 1802]) that is using sprawling forelimb posture and kinematics. A. A forearm and manus forcibly unpronated (0°) to demonstrate the erroneous starting point of dinosaurian pronation according to Bonnan's (2003) radial hypothesis; this dislocation would orient the manual digits laterally in a sprawling forelimb. B. A naturally (for A. mississippiensis) semi-pronated (90°) forearm and manus, which orients the manual digits anteriorly in a sprawling forelimb. C. A forcibly fully pronated (180°) forearm and manus; this dislocation would orient the manual digits medially in a sprawling forelimb. In this and the following figure the upper row of boxes shows the radius and ulna in proximal view, while the lower row of boxes shows their positions in distal view. The orientations represented serve to demonstrate why stem tetrapods with laterally-directed forelimbs are assumed to have evolved semi-pronated forearm morphology from an unpronated morphology, in order to pre-axially rotate the wrist and finger joints 90° so that these joints could participate in locomotion via posteriorly directed flexion (Hutson 2010). Note also that, as a consequence of being a semi-pronated tetrapod, if an A. mississippiensis inturns its elbows to the body wall, then the manual digits will then point laterally (Vialleton 1924). Bonnan's (2003) radial hypothesis assumes that condition A was the starting point of quadrupedal dinosaur evolution instead of condition B; see text for further discussion. This and all subsequently figured A. mississippiensis forelimb elements are from the left forelimb of a juvenile specimen (FMNH 284695).

Fig. 3 in Quadrupedal dinosaurs did not evolve fully pronated forearms: New evidence from the ulna

Fig. 3. The grades of forearm pronation, the incidence of tubular manual cross sections, and the prevalence of proximal radial migration overlaid onto a cladogram of major tetrapod clades. Note the isolated convergence upon tubular manual cross sections in non-avian saurischian and ornithischian dinosaurs. See Vialleton (1924) for reports of proximal radial migration in tetrapods. See text for a discussion of the reasoning that dinosaurs retained semi-pronation. Cladogram after Gauthier (1986).

Fig. 1 in Quadrupedal dinosaurs did not evolve fully pronated forearms: New evidence from the ulna

Fig. 1. An example of the stylized tetrapod skeleton and terminologies that are traditionally used to demonstrate the grades of forearm pronation. A. The three anatomical planes. B. Simplified tetrapod limbs with uniplanar joint alignments in the transverse plane, showing general limb terminology, including the directions required to pronate/supinate the limb segments, and thereby joint planes of action distal to these segments. Note that, like the hindlimb in this traditional characterization, the three major tetrapod forelimb bones are assumed to plesiomorphically lack any diaphyseal torsion or oblique planes of joint flexion/extension.

Fig. 5 in Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain

Fig. 5. Bivariate analysis comparing height (in mm) against posterior denticles per millimeter,?Dromaeosauridae,?Pyroraptor olympius, and?Richardoestesia from the South Pyrenees area are compared against a sample of Dromaeosaurus, Saurornitholestes, Richardoestesia, and Troodon from the collections of the Royal Tyrrell Museum of Palaeontology, the Richardoestesia-like tooth from the site of Suterranya, Catalonia, Spain (Prieto-Márquez et al. 2000) and Pyroraptor olympius Allain and Taquet, 2000 from Provence (Ronan Allain, personal communication 2013).

Fig. 8 in Quadrupedal dinosaurs did not evolve fully pronated forearms: New evidence from the ulna

Fig. 8. Comparison of standardized views of semi-pronated forearms in ornithischian dinosaurs that utilized quadrupedalism. A. A tracing of a representative neosauropod sauropodomorph, Apatosaurus excelsus (Marsh, 1879) (YPM 1980), in distal view. B. A hadrosaur, Edmontosaurus annectens (Marsh, 1892) (USNM 3814), reversed, in distal view. C. A stegosaur, Stegosaurus sp. (USNM 11659), reversed, in flexor (C1), distal (C2), and pre-axial (C3) views. D. A ceratopsid, Torosaurus cf. latus Marsh, 1891 (high fidelity YPM 57489 cast of MPM VP6841), reversed, in flexor (D1), distal (D2), and pre-axial (D3) views. E. An ankylosaur, Sauropelta edwardsorum Ostrom, 1970 (AMNH 3032), in flexor (E1), distal (E2), and pre-axial (E3) views. Pre-axial views are in full extension and scaled to equal radial length. Distal views not to scale.

Fig. 4 in Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain

Fig. 4. Principal component analysis of the South Pyrenees Basin sample and the Royal Tyrrell Museum of Palaeontology sample; a chart displaying two first principal components, PC1 and PC2.

Fig. 1. A in Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain

Fig. 1. A. Locations of the palaeontological sites of Laño, Vicari 4, Montrebei, Fontllonga 6, Figuerola 2, and Blasi. B. Correlation of the uppermost Cretaceous and lowermost Tertiary deposits in the southern Pyrenees, showing the stratigraphic levels of the studied localities. MPU, Mid-Paleocene unconformity; S1, S2, depositional sequences (Robador 2005).

Fig. 6 in Quadrupedal dinosaurs did not evolve fully pronated forearms: New evidence from the ulna

Fig. 6. Comparison of true standardized views of forearm pronation in tetrapods that have evolved a fully pronated manus, in flexor (A1–C1) and distal (A2–C2) views. A. A representative metatherian mammal (marsupial), Virginia opossum Didelphis virginiana Kerr, 1792 (FMNH 166984), oriented in a fully pronated orientation. B. A representative chameleon, Furcifer pardalis (Cuvier, 1829) (FMNH 250433). C. A representative anuran (toad), Bufo blombergi Myers and Funkhouser, 1951 (FMNH 210096). Flexor views are scaled to equal radial length; distal views are not to scale. The reader should note that the distal radial epiphyses of small therians and chameleons are often not located at exactly 180° of pronation relative to their ulnae, as it may not be necessary due to a crouching forelimb posture with moderately abducted elbows (Hutson 2010). Note also that, when present, the full pronation of an anuran manus is accomplished via carpal, not radial torsion (see Schwarz 1935: fig. 10), while the distal radial epiphysis remains in full contact with the plesiomorphically semi-pronated articulation (Ecker and Wiedersheim 1896).

Fig. 1 in Vertebral fusion in two Early Jurassic sauropodomorph dinosaurs from the Lufeng Formation of Yunnan, China

Fig. 1. Location and stratigraphic context of the specimens. A. Location (dinosaur silhouette) of the Dalishu bonebed locality in Yunnan Province, China. B. Stratigraphic section of Lower Jurassic strata in the Lufeng Basin. Based on Xing et al. (2013).

Fig. 5. The 4 in Vertebral fusion in two Early Jurassic sauropodomorph dinosaurs from the Lufeng Formation of Yunnan, China

Fig. 5. The 4th and 5th caudal vertebrae of Sauropoda gen. et sp. indet. (ZLJ 0033) from Dalishu bonebed, Lower Jurassic, in right lateral (A), left lateral (B), dorsal (E), and ventral (F) views; anterior (C, G) and posterior (D, H) views of 4th and 5th caudal vertebrae, respectively. Red line in F shows location of the chevron articulation; red line in G shows boundary between proliferation and centrum.

Fig. 2. The 7 in Vertebral fusion in two Early Jurassic sauropodomorph dinosaurs from the Lufeng Formation of Yunnan, China

Fig. 2. The 7th and 8th cervical vertebrae of spondyloarthropathy dinosaur Lufengosaurus huenei Young, 1941 (ZLJ T001) from Dalishu bonebed, Lower Jurassic, in right lateral (A), left lateral (B), posterior (C), dorsal (D), ventral (E), and anterior (F) views.

Fig. 3 in Vertebral fusion in two Early Jurassic sauropodomorph dinosaurs from the Lufeng Formation of Yunnan, China

Fig. 3. Details of the rugose surfaces of the posterior neural-spine faces of the 7th (A) and 8th (B) cervical vertebra of spondyloarthropathy dinosaur Lufengosaurus huenei Young, 1941 (ZLJ T001) from Dalishu bonebed, Lower Jurassic.

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Fig. 2 in Evidence for a sauropod-like metacarpal configuration in stegosaurian dinosaurs

Fig. 2. Correctly articulated (vertically oriented, based on the results of this study) right manual skeleton of the stegosaur Stegosaurus sulcatus, from the Upper Jurassic of Wyoming, USA USNM 4937, shown in four oblique views with (A–D, I) and without (E–H, J) carpals, and with each metacarpal shown individually in lateral view with its associated phalanx or phalanges correctly articulated (K–N). Roman numerals refer to digit number.

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Fig. 1 in Evidence for a sauropod-like metacarpal configuration in stegosaurian dinosaurs

Fig. 1. Manual skeletal configuration in stegosaurs and other dinosaurs. A. Cladogram of Dinosauria with proximal views of metacarpals, showing convergent evolution of tightly curved metacarpal arc in Sauropoda and Thyreophora; Herrerasaurus after Sereno (1993), sauropods after Bonnan (2003), Camptosaurus after Carpenter and Wilson (2008), and Peloroplites after Carpenter et al. (2008). B. Articulated manus of the sauropod Camarasaurus lentus, CM 11338 (after Gilmore 1925). C. Articulated manus of the stegosaur Stegosaurus armatus, USNM 4934 (right, after Gilmore 1914) as found in situ, showing that the metacarpals form a vertical tube (C. lentus) or semi−tube (S. armatus). D. Articulated metacarpals of the stegosaur Kentrosaurus aethiopicus in proximal view (modified from Hennig 1925). E. Previous, incorrect reconstruction of the manus of USNM 4937 (Gilmore 1914), a stegosaur of indeterminate genus and species (Maidment et al. 2008), in cranial view, showing unnatural gap between carpals (arrow) (likely Stegosaurus sulcatus, Upper Jurassic, USA; see text). F. Previous, incorrect reconstruction of the manus of USNM 4937 (Gilmore 1914) in proximal view, showing unnatural gaps between metacarpals (arrows). G. Incorrectly articulated right hand of the stegosaur USNM 4937 viewed from above. H. Incorrectly articulated right hand of the stegosaur USNM 4937 viewed from obliquely behind and to the left. I. Cranial view of mounted right forelimb of Stegosaurus armatus AMNH 650, showing distal contact between radius and ulna; carpus and manus are cast from a different specimen. Scale bar applies to photographs only; line illustrations not to scale. Roman numerals refer to digit number.

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Fig. 4 in Variation in premaxillary tooth count and a developmental abnormality in a tyrannosaurid dinosaur

Fig. 4. Hypothetical zones of inhibition during premaxillary odontogenesis in tyrannosaurids. The outline represents a tyrannosaurid premaxilla in posteromedial view. A. A normal four−toothed premaxilla of a tyrannosaurid. B. TMP 2007.20.124 with only three premaxillary teeth. A zone of inhibition is a third larger in TMP 2007.20.124 than in a normal tyrannosaurid premaxilla, excluding the fourth tooth position from the alveolar margin.