APPENDIX 1
List of characters and character state used in the phylogenetic analyses. Format follows Sereno (2007). Characters with no citation following are new in this analysis. * indicates an ontogenetically sensitive character, o indicates an ordered character.
1. *Preorbital skull length: less than or equal to 50% of the basal skull length (0); more than 55% of the basal skull length (1). (Modified from Weishampel et al., 2003)
2. Premaxilla, width of the oral margin in dorsal view: narrower than width across orbital region of skull roof (0), equals or exceeds width across orbital region of skull roof (1). (Modified from Norman, 2002)
3. *Premaxilla, shape of subnarial region: narial portion of the body of the premaxilla slopes steeply from the external naris to the oral margin (0); ventral premaxilla flares laterally to form a partial floor of the narial fossa (1). (Weishampel et al., 2003; Butler et al., 2008)
4. Premaxilla, oral margin reflected dorsally: absent (0); present (1). (Modified from Norman, 2002)
5. Premaxilla, oral margin denticulation: absent (0); present (1). (Weishampel et al., 2003; McDonald, 2012a)
6. *Premaxilla, posterolateral process contacts lacrimal: absent (0); present (1). (Modified from Weishampel et al., 2003, Milner and Norman, 1984, 1990; Sereno, 1984, 1986)
7. Premaxilla, posterolateral process, shape: tapers to a point (0); does not taper, caudal end is blunt (1). (modified from McDonald, 2012a, Prieto-Márquez and Salinas, 2010).
8. Premaxilla, position of the ventral (oral) margin: level with the maxillary tooth row (0); ventral to maxillary tooth row (1). (Norman, 2002; Butler et al., 2008)
9. Premaxilla, medial dorsal (nasal) process contacts the nasal: present (0); absent (1). (Butler et al., 2008)
10. Premaxilla, discrete rugose rostral margin, extending onto the medial dorsal (nasal) process: absent (0); present (1).
11. External naris, size: small, entirely overlies the premaxilla (0); enlarged, extends posteriorly to overlie both the premaxilla and maxilla (1). (Weishampel et al., 2003; Butler et al., 2008)
12. Nasals, deep elliptic fossa along midline: absent (0); present (1). (Butler et al., 2008)
13. *Nasals, placement of most dorsal point: caudally, at junction with frontals (0); more rostral, such that the nasal extends further dorsally than the frontals (1).
14. Nasals, thickened, paired domes on caudal portion: absent (0); present (1).
15. Premaxilla-maxilla boundary, fossa or foramen just dorsal to the oral margin: absent (0); present (1). (Butler et al., 2008)
16. Maxilla, diastema: absent, maxillary teeth begin directly caudal to the premaxilla (0); present, first maxillary tooth is at least one crown width away from the premaxilla (1). (Modified from Butler et al., 2008)
17. Maxilla, diastema length: one to two tooth widths (0); three tooth widths or more (1).
18. Maxilla, rostrolateral process in addition to the premaxillary process: absent (0); present (1). (Weishampel et al., 2003; Butler et al., 2008)
19. Maxilla, rostrolateral process of the maxilla, length: shorter than the premaxillary process (0); longer than the premaxillary process (1).
20. Maxilla, prominent rostrolateral boss articulates with the lateral face of the premaxilla: absent (0); present (1). (Butler et al., 2008)
21. Maxilla, dorsal process, shape in lateral view: narrow, parallel-sided, (0); broad and triangular (1). (Modified from Norman, 2002)
22. o*Maxilla, dorsal process, position: rostral to the midpoint of the maxilla (0); at the midpoint (1); caudal to the midpoint (2).
23. Maxilla, emargination of the caudal end of the tooth row: absent (0); present (1). (new)
24. Maxilla, form of emargination: smooth curve (0); angular ridge (1).
25. Maxilla, horizontal ridge, caudal portion covered by a series of obliquely inclined ridges: absent (0); present (1). (Modified from Boyd et al., 2009)
26. Maxilla, shape of tooth row in ventral view: medially bowed, with rostral and caudal ends curving laterally (0); laterally bowed (1); straight (2). (McDonald, 2012a)
27. Maxilla, articulation with lacrimal: simple scarf or butt joint (0); lacrimal fits into slot between medial and lateral portions of the dorsal process of the maxilla (1). (Modified from Butler et al., 2008)
28. Maxilla, exclusion from the antorbital fenestra by rostral extension of the jugal: absent (0); present (1).
29. Antorbital fenestra: present (0); absent (1). (Butler et al., 2008)
30. Antorbital fenestra, shape: triangular (0); oval or circular (1). (Butler et al., 2008)
31. Antorbital fenestra, size: large (at least 15% basal skull length) (0); relatively small (10% basal skull length or less) (1); reduced to a small foramen (2). (Modified from Weishampel et al., 2003)
32. Antorbital fossa, additional fenestra(e) within fossa rostral to external antorbital fenestra: absent (0); present (1). (Butler et al., 2008)
33. Lacrimal-nasal contact: present (0); absent (1). (Norman, 2002)
34. Orbit shape: circular (0), subrectangular at least in its lower margin (1); dorsoventrally elongated, either oblong or subtriangular (2). (Modified from Winkler et al., 1997)
35. *Frontal, proportions: short and broad (length roughly equal to width) (0); narrow and elongate (length roughly 2 times the width) (1); very elongate (length roughly 3 times the width) (2). (Modified from Weishampel et al., 2003; Butler et al., 2008; state 2 based on Barrett and Han, 2009)
36. Frontal, shape in dorsal view: widest point is posterior to midorbit level (0); widest point is at midorbital level (1). (Modified from Boyd et al., 2009)
37. Frontal, participation in dorsal orbital rim: present (0); absent (1). (Norman, 2002; McDonald, 2012).
38. Frontal, participation in the border of the supratemporal fenestra: present (0); absent, excluded by contact between the postorbital and parietal (1).
39. Palpebral(s): free and separated from orbital rim, articulate only to the prefrontal (0); fused to orbit rim (1). (Modified from Butler et al., 2008)
40. Palpebral(s), shape in dorsal view: rod-shaped (0); platelike (1). (Butler et al., 2008)
41. Palpebral/supraorbital, number of ossified segments: one (0); two (1). (Butler et al., 2008)
42. *Palpebral(s), length relative to rostrocaudal width of orbit if unfused to orbital rim: does not traverse entire width of orbit (0); traverses entire width of orbit (1). (Butler et al. 2008)
43. Palpebral, dorsoventrally flattened and rugose along the medial and distal edges: absent (0); present (1). (Boyd et al., 2009)
44. *Postorbital, projection into orbital margin or rugose area for articulation of the palpebral: absent (0); present (1). (Modified from Butler et al., 2008)
45. Postorbital, shape of squamosal process: tapered (0); bifurcated (1). (modified from McDonald, 2012; Prieto-Márquez, 2010).
46. *Jugal, proportions: length of rostral process greater than or equal to caudal process (0); caudal process longer than rostral process (1).
47. Jugal, exclusion of the rostral process from the antorbital fenestra by lacrimal-maxilla contact: absent (0); present (1). (Weishampel et al., 2003; Butler et al., 2008)
48. o Jugal, shape of rostral end in lateral view: tapering (0); near parallel sided with a rounded or blunt rostral end (1) expanded dorsoventrally (2). (Modified from Norman, 2002)
49. o Jugal, type of suture with maxilla: scarf joint (0); 'finger-in-recess' joint, (1); butt-joint, (2). (Norman, 2002).
50. Jugal, boss extending laterally from suborbital portion: absent (0); present (1). (Butler et al., 2008)
51. Jugal, articulation with ectopterygoid: present (0); absent (1). (Head, 1998; Norman, 2002; McDonald, 2012)
52. Jugal, caudal ramus, bifurcated caudal end: absent (0); present (1). (Butler et al., 2008)
53. Jugal, participation in infratemporal fenestra: forms cranial and ventral margin (0); also forms part of caudal margin (1). (Weishampel et al., 2003; Butler et al., 2008)
54. Infratemporal fenestra: ventral end narrower than dorsal end: absent (0); present (1).
55. Jugal, form of articulation with quadratojugal: contact is relatively simple, consisting of a butt or high-angle scarf joint, jugal lies entirely lateral to the quadratojugal (0); tongue and-groove contact whereby the jugal lies lateral to the quadratojugal dorsally while ventrally the jugal lies medial to the quadratojugal (1). (Weishampel et al., 2003)
56. Jugal, shape of ventral edge: straight (0); sinuous (1).
57. Quadratojugal, shape: a central body with a dorsally projecting quadrate process that meets or nearly meets the squamosal (often on the medial side of the quadrate) (0); no dorsal process, element is small and blocky (1).
58. Quadratojugal, foramen through center of element: absent (0); present (1).
59. Quadratojugal foramen, position: located within quadratojugal (0); on jugal-quadratojugal boundary (1). (Modified from Butler et al., 2008)
60. Quadratojugal, position of ventral margin relative to mandibular condyle of the quadrate: approaches the mandibular condyle (0); well-removed from the mandibular condyle (1). (Butler et al., 2008)
61. *Quadrate, ventral portion of shaft: cranially convex (0); straight (1); caudally convex (2). (Modified from Butler et al., 2008)
62. Quadrate, caudally curved proximal portion: present (0); absent, straight (1). (Boyd et al., 2009)
63. Quadrate buttress ("hamular process"): absent (0); present (1).
64. Quadrate, prominent oval fossa on pterygoid ramus: absent (0); present (1). (Butler et al., 2008)
65. Quadrate (paraquadratic) foramen or notch on boundary between quadrate and quadratojugal: absent (0); present (1). (Butler et al., 2008; Norman, 2002)
66. *Quadrate, paraquadratic foramen or notch, size: small, height less than 1/8th quadrate height (0); large, height more than 1/7th quadrate height (1). (Modified from Butler et al., 2008)
67. Quadrate, paraquadratic notch completely covered by quadratojugal: absent (0); present (1).
68. Quadrate, articular condyle shape: much wider transversely than craniocaudally (0); transverse width less than or equal to craniocaudal length (1). (Modified from Norman, 2002)
69. o Quadrate, mandibular condyle proportions: medial condyle is larger than lateral condyle (0); quadrate condyles subequal in size (1); lateral condyle is larger than medial (2). (Butler et al., 2008)
70. o Quadrate, mandibular condyles, orientation of distal margin relative to long axis of quadrate: dorsomedially sloped (0); horizontal (1); dorsolaterally sloped (2).
71. Squamosal, ventral (quadratic) process length: less than 25% length of the quadrate (0); greater than 30% (1). (Modified from Boyd et al., 2009)
72. Squamosal, relative lengths of pre- and post-quadratic processes: pre-quadratic process is longer than or subequal to the post-quadratic (0); pre-quadratic process is shorter (1).
73. o Squamosal, relationship of right and left squamosals on skull roof: widely separated by parietal (0); separated by only a narrow band of the parietal (1); in broad contact with each other (2) (Horner et al., 2004; McDonald, 2012)
74. Paroccipital processes, shape: extend laterally (0); pendant, distal end curves ventrally (1). (Modified from Weishampel et al., 2003; Butler et al., 2008)
75. Posttemporal foramen/fossa, position: totally enclosed with the paroccipital process (0); forms a notch in the dorsal margin of the paroccipital process, enclosed dorsally by the squamosal (1); enclosed entirely by the squamosal (2). (Modified from Butler et al., 2008)
76. Supraoccipital, participation in the dorsal margin of the foramen magnum: present (0); absent, excluded by exoccipitals, (1).
77. *Supraoccipital, sharp, well-defined median nuchal crest: absent (0); present (1).
78. Supraoccipital, inclination of caudal surface: rostrodorsally inclined (0); vertical (1) (McDonald, 2012; modified from Horner et al., 2004)
79. Basioccipital: participation in the ventral border of the foramen magnum: present (0); absent, excluded by the exoccipitals (1).
80. Basioccipital, ventral keel: absent (0); present (1). (Boyd et al., 2009)
81. o Basipterygoid processes, orientation: anteroventral (0); ventral (1); posteroventral (2). (Butler et al., 2008)
82. Premaxilla, contact with vomer: present (0); absent, excluded by maxillae (1). (Modified from Butler et al., 2008)
83. Pterygoid, contact with maxilla at posterior end of tooth row: absent (0); present (1). (Butler et al., 2008)
84. Predentary, size: short, caudal oral margin of the premaxilla opposes rostral dentary margin (or teeth) (0); subequal in length to the oral margin of the premaxilla (1). (Modified from Butler et al., 2008)
85. Predentary, rostral end in dorsal view: rounded (0); pointed (1); straight transverse margin (2). (Weishampel et al., 2003; Butler et al., 2008)
86. Predentary, sulci extending rostrally from the intersections of the lateral processes and the ventral process: absent (0); present (1). (Modified from McDonald, 2012a)
87. Predentary, denticulate oral margin: absent (0); present (1). (Weishampel et al., 2003; Butler et al., 2008)
88. Predentary, ventral process: single (0); bilobate (1). (Butler et al., 2008)
89. Predentary, ventral process, depth of bifurcation: restricted to distal end (0); deep bifurcation (1).
90. Predentary, short midline process dorsal to dentary symphysis: absent (0); present (1). (Modified from McDonald, 2012)
91. o Predentary, relative length of the dorsal and ventral processes: ventral process longer than dorsal (0); processes roughly equal in length (1); dorsal process longer than ventral (2).
92. Dentary, diastema between predentary and first tooth: absent (0); present (1).
93. Dentary, length of diastema: short, the width of one to two teeth (0); long, the width of three teeth or more (1). (Modified from Norman, 2002)
94. Dentary, rostral extent of Meckel's groove meets the dentary symphysis: present (0); absent, ends more caudally (1).
95. *Dentary ramus, shape of rostral end: straight (0); strongly downturned (1). (Norman, 2002)
96. o Dentary, position of rostral tip: dorsal to the ventral margin of the dentary ramus (0); level with the ventral border of the dentary ramus (1); well below the ventral border of the dentary ramus (2).
97. o*Dentary, relationship of dorsal and ventral margins (under the tooth row): converge anteriorly (0); subparallel (1); diverge anteriorly (2). (Modified from Butler et al., 2008; Norman, 2002; Weishampel et al., 2003; McDonald, 2012a)
98. Dentary, shape of tooth row in dorsal view: straight (0); bowed medially (1).
99. Dentary, position of apex of curve in medially bowed tooth row: at mid-length (0); in the caudal half of the dentary (1). (McDonald, 2012a, modified from Prieto-Márquez et al., 2006)
100. Dentary, caudoventral extension below angular: absent (0); present (1).
101. Dentary, coronoid process, height; short, extends no more than one tooth height above tooth row (0); distinctly higher (1). (Modified from Weishampel et al., 2003)
102. Dentary, medial edge of coronoid process, position: in line with dentition (0); lateral to dentition (1). (Modified from Butler et al., 2008)
103. o*Dentary: caudal extent of tooth row: terminates rostral to coronoid process (0); terminates between rostral margin and apex of coronoid process (1); terminates directly ventral to apex or more caudally (2). (Modified from Wu and Godefroit, 2012)
104. Dentary: if coronoid is lateral to dentition, trough present between toothrow and coronoid: absent (0) present (1).
105. o Dentary, coronoid process orientation: caudally inclined (0); subvertical (1); rostrally inclined (2). (modified from Prieto-Márquez et al., 2006; McDonald, 2012a)
106. Coronoid process, shape in lateral view: subtriangular, tapers dorsally (0); rectangular (rostral and caudal margins are nearly parallel (1).
107. Coronoid process, rostrocaudally expanded apex: absent (0); present (1). (Modified from McDonald, 2012a)
108. Coronoid process, surangular contribution: present (0); absent (1).
109. o Coronoid process, relative craniocaudal widths of dentary and surangular (at midpoint of coronoid process): surangular wider than dentary (0); subequal (1); surangular narrower than dentary (2).
110. External mandibular fenestra, situated on dentary-surangular-angular boundary: present (0); absent (1). (Butler et al., 2008)
111. Surangular, small fenestra positioned dorsally on or near the dentary joint: absent (0); present (1). (Modified from Butler et al., 2008)
112. Surangular, position of small rostral fenestra: lies on boundary of dentary and surangular (0); lies within the surangular (1).
113. Surangular foramen near mandibular glenoid: present (0); absent (1). (Norman, 2002)
114. Surangular foramen, position relative to the lateral lip of the glenoid: foramen is directly ventral to process (0); foramen is placed more rostrally (1).
115. Surangular, lateral lip of the glenoid expanded dorsally into a distinct process: absent (0); present (1). (Modified from Butler et al., 2008; Boyd et al., 2009)
116. Angular position, visible on the lateral surface of the mandible: present (0); absent (1). (Norman 2002)
117. Jaw joint, position: weakly depressed ventral to toothrow (0); strongly depressed ventrally, more than 30% of the height of the quadrate is below the level of the maxilla (1). (Modified from Butler et al., 2008)
118. Articular, retroarticular process: elongate (0); rudimentary or absent (1). (Butler et al., 2008)
119. Premaxillary teeth: present (0); absent (1). (Butler et al., 2008)
120. o* Premaxillary teeth, number: six (0); five (1); two (2); one (3). (modified from Butler et al., 2008)
121. Premaxillary teeth, mesiodistal crown expansion above root: absent (0); present (1). (Butler et al., 2008)
122. Premaxillary teeth, spacing: closely spaced (0); gaps of about one crown width between teeth (1).
123. o*Maxillary teeth, number; 13 or fewer tooth positions (0); 14 to 16 positions (1); 18-28 positions (2); 30 or more positions (3). (Modified from Weishampel et al., 2002)
124. o*Dentary teeth, number. 13 or fewer tooth positions (0); 14 to 16 positions (1) 18 to 25 positions (2); 27 or more positions (3). (Modified from Weishampel et al., 2002)
125. Cheek teeth; dentary teeth extend farther rostrally than maxillary teeth: absent (0); present (1).
126. Cheek teeth, spaces between the roots: present (0); absent, teeth are closely packed (1).
127. o Maxillary teeth, morphology of roots: distinct neck between crown and root (0); crown tapers to root (1); crown not significantly wider than root (2). (Modified from Boyd et al., 2009).
128. Dentary teeth, morphology of roots: distinct neck between crown and root (0); crown tapers to root (1). (Modified from Boyd et al., 2009)
129. o*Maxillary teeth, ratio of crown height to width (for unworn teeth): less than 1.2 (0); 1.25 to 1.9 (1); greater than 2 (2).
130. o*Dentary teeth, ratio of crown height to width (for unworn teeth): less than 1.5 (0); 1.7 to 2.2 (1); greater than 2.5 (2).
131. Cheek teeth: number of wear facets on each tooth: two (0); one (1).
132. Cheek teeth, apicobasally extending thickening of crown along midline (not a primary ridge): absent (0); present (1).
133. Cheek teeth, form of marginal denticles: simple, arcuate (0); labiolingually expanded, with a mammillated edge (1); absent or reduced to small papillae (2). (Modified from Norman, 2002)
134. Cheek teeth, enamel distribution: symmetrical (0); asymmetrical: thicker on the labial side of maxillary teeth and on the lingual side of dentary teeth (1). (Weishampel et al. 2003; Butler et al., 2008)
135. Cheek teeth, apicobasally extending ridges on labial side of maxillary teeth and lingual side of dentary teeth: absent (0); present (1). (Butler et al., 2008)
136. o Maxillary teeth, maximum number of ridges extending from the base to the tip of the crown on labial side of teeth: primary ridge only (0); one primary and one secondary ridge (1); three to eight ridges (2); ten to twelve ridges (3); fourteen to seventeen ridges (4).
137. o Dentary teeth, maximum number of ridges extending from the base to the tip of the crown on lingual side of teeth: primary ridge only (0); two to four ridges (1); five to eight ridges (2); nine to eleven ridges (3); twelve to seventeen ridges (4).
138. Cheek teeth: apicobasally extending ridges on cutting surface of unworn teeth (lingual surface of maxillary teeth, labial surface of dentary teeth): absent (0); present (1).
139. Maxillary teeth, primary ridge on labial side of crown: absent (ridges may be present, but none is more prominent than others) (0); present (1). (Modified from Weishampel et al., 2003; Butler et al., 2008)
140. Maxillary teeth, primary ridge size: only slightly larger than secondary ridges (0); much larger than secondary ridges (1). (Modified from Weishampel et al., 2003 and Butler et al., 2008)
141. Dentary teeth, primary ridge on lingual side of crown: absent (ridges may be present, but none is more prominent than others) (0); present (1). (Modified from Weishampel et al. 2003; Butler et al., 2008)
142. Dentary teeth, primary ridge size: only slightly larger than secondary ridges (0); much larger than secondary ridges (1). (Modified from Weishampel et al., 2003 and Butler et al., 2008)
143. Maxillary teeth, primary ridge position: centered, although some teeth within the same dental battery may display a slight offset of the primary ridge (0); offset, giving crown asymmetrical appearance (1). (Modified from Butler et al., 2008 and Prieto-Márquez and Salinas., 2010)
144. Dentary teeth, primary ridge position: centered, although some teeth within the same dental battery may display a slight offset of the primary ridge (0); offset, giving crown asymmetrical appearance (1). (Modified from Butler et al., 2008; Prieto-Márquez and Salinas., 2010)
145. Cheek teeth, secondary ridges, shape in cross-section: thick and rounded, composed of both enamel and dentine (0); thin and sharp-edged, formed by crenulations in enamel (1).
146. Maxillary teeth, base of crown defined by an everted lip which makes the crown slightly inset from the root: absent (0); present (1).
147. Dentary teeth, base of crown defined by an everted lip which makes the crown slightly inset from the root: absent (0); present (1).
148. Cheek teeth, at least moderately developed labiolingual expansion at the base of the crown ("cingulum"): present (0); absent (1). (Butler et al., 2008)
149. Maxillary and dentary crowns, relative width: maxillary crowns approximately equal in width with dentary crowns (0); maxillary crowns narrower (1). (Norman, 2002)
150. o Dentary teeth, maximum number of functional teeth exposed on the occlusal plane: one (0); one functional tooth rostrally and caudally, and up to two teeth at and approaching the middle of the dental battery (1); three functional teeth throughout most of the dental battery, gradually decreasing to two near the rostral and caudal ends of the dentary (2). (Prieto-Márquez and Salinas, 2010)
151. o Dentary teeth, maximum number of replacement teeth: one (0); two (1); three or more (2). (Norman, 2002)
152. Dentary alveoli: distinct, separate alveoli (0); parallel grooves lining a continuous dental battery (1). (Modified from Wu and Godefroit, 2012)
153. o Cervical vertebrae, number: nine or fewer cervical vertebrae (0); 10 to 11 (1); 12 or more (2). (Modified from Butler et al., 2008, Weishampel et al., 2003 and Prieto-Márquez and Salinas., 2010)
154. Axis, neural spine, shape of dorsal margin in lateral view: concave to straight (0); convex (1); sinuous (convex in the cranial portion and concave in the caudal portion) (2). (last state adapted from Prieto-Márquez and Salinas., 2010)
155. Axis, postzygapophyses, position: below dorsal margin of the neural spine (0); at or above the dorsal margin of the neural spine (1).
156. Postaxial cervical vertebrae, epipophyses: present on anterior vertebrae (0); absent (1). (Butler et al., 2008)
157. *Postaxial cervical vertebrae, form of central surfaces: amphicoelous (0); at least slightly opisthocoelous (1). (Butler et al., 2008)
158. Postaxial cervical vertebrae, degree of opisthocoely: cranial surfaces slightly convex (0); cranial surfaces distinctly convex (1).
159. Postaxial cervical vertebrae, proportions of centra: craniocaudal length about equal to dorsoventral height (0); very elongate: craniocaudal length more than twice the dorsoventral height (1).
160. Cervical vertebrae, ventral keel on centra: absent (0); present (1).
161. Dorsal vertebrae, number: 15 or fewer dorsal vertebrae (0); 16 or more dorsals (1). (Weishampel et al., 2003; Butler et al., 2008)
162. o Dorsal vertebrae, length of mid-dorsal neural spines: short and rectangular, height and length roughly equal (0); height more than twice length (1); height more than four times length (2). (Norman, 2002)
163. Dorsal vertebrae, cranial and mid dorsals, ventral keel on centra: absent (0); present (1).
164. Dorsal vertebrae, proportions of mid to posterior centra: craniocaudal length is subequal to or longer than dorsoventral height (0); length is much shorter than height (1).
165. Dorsal vertebrae, shape of neural spines in posterior dorsals: straight (0); bowed caudally (1).
166. Dorsal vertebrae, prezygapophyses on posterior dorsals extend well past the cranial edge of the centrum: absent (0); present (1).
167. Dorsal vertebrae, posterior dorsals, length of transverse processes: less than or equal to centrum height (0); greater than centrum height (1).
168. Ossified intercostal plates: absent (0); present (1).
169. Sternal segments of the anterior dorsal ribs: not ossified (0); at least partially ossified (1). (Modified from Weishampel et al 2003)
170. Sacral vertebrae, number, including dorsosacrals and sacrocaudals: five or fewer (0); six or more (1). (Weishampel et al., 2003; Butler et al., 2008)
171. Sacral vertebrae, transverse thickening and increase in rugosity at tips of neural spines: present (0); absent (1).
172. o Sacral vertebrae, neural spines, extent of fusion: spines do not touch (0); some spines overlap, but are not fused (1); spines are fused into a single plate (2).
173. Sacral vertebrae, form of ventral surface of centra: rounded (0); keeled (1); grooved (2).
174. Sacral ribs, coalesce laterally to form a sacral yolk: absent (0); present (1).
175. Caudal vertebrae, proportions of proximal centra: craniocaudal length is subequal to or longer than dorsoventral height (0); length is much shorter than height (1).
176. Caudal vertebrae, ventral sulcus on centra: absent (0); present (1).
177. Caudal vertebrae, position of the most distal caudal rib (transverse process): 12th caudal vertebrae or more proximal (0); 13th or more distal (1).
178. Caudal vertebrae, height of neural spines on proximal caudals: height the same or up to 50% taller than the centrum (0); more than 50% taller than the centrum (1). (Weishampel et al., 2003; Butler et al., 2008)
179. Caudal vertebrae, orientation of neural spines on proximal caudals: project dorsally from centra (0); project caudodorsally from centra (1).
180. Caudal vertebrae, shape of neural spines on proximal caudals: straight (0); bowed caudally (1).
181. Caudal vertebrae, length of transverse processes on proximal caudals: longer than or subequal to neural spine height (0); shorter than neural spine (1). (Modified from Butler et al., 2008)
182. Caudal vertebrae, craniocaudal length of the distal facet for chevrons on proximal caudals: shorter than the ventral surface of the vertebrae (between the proximal and distal facets) (0); longer than the ventral surface (1).
183. Caudal vertebrae, proportions of articular facets for chevrons: the distal facet on a given vertebra is much larger than the proximal facet (0); facets are subequal (1).
184. Caudal vertebrae, form of chevron articular facets: fairly flat surface (0); deep fossa surrounded by a rim (1).
185. Caudal vertebrae, length of prezygopophyses on distal caudals: extend slightly over the preceding vertebra (0); elongate, extend nearly to the midpoint of the preceding vertebra (1).
186. o Chevrons, position of most proximal chevron: distal end of first caudal vertebra (0); distal end of second caudal vertebra (1); distal end of third caudal vertebra or more distal (2).
187. Chevrons, length relative to the length of the neural spines in the proximal caudal vertebrae: chevrons shorter than or equal in length to the neural spines, (0); chevrons longer than neural spines (1). (Modified from Prieto-Márquez and Salinas., 2010)
188. Chevrons, shape: edges are near parallel in lateral view, often with slight distal expansion (0); strongly asymmetrically expanded distally, width greater than length in mid caudals, (1). (Modified from Butler et Al., 2008)
189. Sternal plates, rod-like caudolateral process: absent (0); present (1). (Modified from Butler et al., 2008, Norman, 2002)
190. Sternals, caudolateral process, shape of cross-section: round (0); flattened (1).
191. Sternal, caudolateral process, length relative to that of the craniomedial plate: caudolateral process slightly shorter or as long as the craniomedial plate (0); caudolateral process longer than the craniomedial plate (1). (Prieto-Márquez and Salinas., 2010).
192. Sternal, pronounced caudomedial process ("posterior apron" of Norman, 2015) projects from plate of sternal in addition to caudolateral process: absent (0); present (1).
193. Sternals, midline fusion: absent (0); present (1).
194. *Forelimb, proportions of humerus and scapula: scapula longer or subequal to the humerus (0); humerus substantially longer than the scapula (1). (Weishampel et al., 2003; Butler et al., 2008)
195. Scapula, acromion process size: weakly developed (0); projects prominently from the cranio-dorsal edge of the proximal scapula (1). (Modified from Butler et al., 2008)
196. Scapula, acromion process, length: does not extend beyond the edge of the coracoid (0); extends past the coracoid (1).
197. Scapula, deltoid ridge shape: wide and rounded (0); sharp, narrow (1).
198. *Scapula, deltoid ridge, orientation relative to long axis of scapula: close to parallel (0); more than 20 degrees from axis (1).
199. Scapula, supraglenoid fossa: absent (0); present (1).
200. o*Scapula, length of blade relative to minimum width: short and broad, ratio of length to width 5.7 or less (0); ratio of length to width 6 to 7.3 (1); elongate, ratio of 7.5 or greater (2). (modified from Butler et al., 2008)
201. Scapula blade, shape: expanded distally (0); parallel-sided (1). (Modified from Butler et al., 2008)
202. Scapula, expanded distal blade shape: caudoventral edge curves away from a fairly straight craniodorsal edge, creating a deep asymmetric expansion (0); edges gradually diverge from each other, forming a symmetrically expanded end (1).
203. Scapula blade, shape of dorsal edge in lateral view: straight (0); curved (1). (Norman, 2002)
204. o Scapula and coracoid, relative contributions to the glenoid fossa in transverse width: scapula wider (0); subequal (1); coracoid wider (2).
205. o Scapula and coracoid, relative contributions to the glenoid fossa in craniocaudal length: scapular portion longer (0); subequal (1); coracoid portion longer (2).
206. Coracoid, depth of preglenoid embayment: wide and shallow, depth of embayment less than 40% of its width (0); deep, depth 45% of its width or greater (1).
207. Coracoid, ratio between the length of the scapular articulation and the length of the lateral margin of the glenoid in lateral view: greater than 1.30 (0); less than 1.25 (1).
208. Coracoid, position of lateral opening of coracoid foramen: within coracoid (0); on suture between the coracoid and scapula (1).
209. Coracoid, position of medial opening of coracoid foramen: within coracoid (0); on suture between the coracoid and scapula (1).
210. Coracoid, lateral protrusion on dorsal border for origin of M. biceps: absent (0); present (1).
211. Humerus, length relative to femur: less than 62% of femoral length; (0); greater than 65% of femoral length (1). (Modified from Butler et al., 2008)
212. *Humerus, deltopectoral crest form: a well-developed projection (0); slight projection (1). (Modified from Butler et al., 2008; Norman, 2002)
213. Humerus, deltopectoral crest orientation: projects laterally (0); curves cranially (1).
214. *Humerus, deltopectoral crest length: 40% of total humeral length or less (0); 43% or more (1).
215. Humerus, humeral head position: centered within the proximal end (0); offset towards the medial edge (1).
216. o Humerus, relative width of medial and lateral distal condyles: lateral condyle wider (0); subequal (1); medial condyle wider (2).
217. Ulna, length relative to dorsoventral thickness at mid-shaft: less than 9 (0); greater than 9.5 (1). (Modified from Prieto-Márquez and Salinas., 2010)
218. o Ulna, olecranon process length as a percentage of total ulnar length: 9% or less (0); 9.5 to 15% (1): 17% or greater (2).
219. Ulna, flange on proximal end that wraps around the lateral edge of the radius: absent (0); present (1).
220. Radius, length: less than 67% of the length of the humerus (0); greater than 70% of humeral length (1). (Modified from Norman, 2002)
221. Radius, proportions: relatively gracile, minimal radial width 11% of radial length or less (0); relatively robust, the minimal radial width is equal to or greater than 12% radial length (1). (modified from Weishampel et al., 2003)
222. Radius, notch in proximal view, rostrolateral to articulation with the ulna: absent (0); present (1).
223. Radius, tubercle near proximal end of radius for insertion of M. biceps: absent (0); present (1).
224. Radius, shape in distal view: round to oblong (0); triangular (1).
225. Carpals, full ossification of all elements: present (0); absent (1). (Norman, 2002)
226. Carpals, relative sizes of proximal carpals: radiale is the largest element (0); ulnare is the largest element (1).
227. Carpals, fusion: absent (0); present (1). Modified from (Sereno, 1986; Norman, 1986, 1990, Weishampel et al., 2003)
228. o Carpals, degree of fusion: metacarpal I fused to radiale (0); radiale is further fused to intermedium (1); all carpals and the first metacarpal are fused (2). Modified from (Sereno 1986; Norman 1986, 1990, Weishampel et al., 2003)
229. Carpals, ossified ligaments around carpals: absent (0); present (1).
230. Metacarpal III, ratio between length and width at mid-shaft: short, ratio less than 5 (0); long and slender, ratio greater than 5.5 (1). (modified from Horner et al., 2004; Prieto-Márquez and Salinas., 2010)
231. Manus digit I: present (0); absent (1). (Norman, 2002)
232. Manus digit I, orientation: nearly parallel to the long axis of the antebrachium (0); diverges at least 45 degrees from the antebrachial axis (1). (Modified from Weishampel et al., 2002)
233. Metacarpal I, shape: regular, elongate metacarpal (0); short, block-like (1). (Modified from Norman, 2002)
234. Metacarpals II-IV arrangement: distally divergent (0); nearly parallel (1). (Modified from Norman, 2002)
235. Metacarpals II and IV, relative length: subequal (0); metacarpal II is much shorter than metacarpal IV (1); metacarpal IV is much shorter than metacarpal II (2).
236. Manus digit III, number of phalanges: four (0); three or fewer (1). (Butler et al., 2008)
237. Manus, digit V: present (0); absent (1).
238. Manus, digit V, number of phalanges: one or two phalanges (0); three or four phalanges (1). (Modified from Wu and Godefroit, 2012)
239. Phalanges of manual digits II-IV, length: first phalanx less than twice the length of the second phalanx (0); first phalanx more than twice the length of the second phalanx (1). (Modified from Butler et al., 2008 and Weishampel et al., 2002)
240. Phalanges, extensor fossae on the dorsal surface of the distal end of metacarpals and manual phalanges: absent or poorly developed (0); deep, well-developed (1). (Butler et al., 2008)
241. Manus ungual I, shape: curved and transversely compressed (0); subconical (1). (Sereno, 1984, 1986; Norman, 1986, 1990, 2002, Weishampel et al., 2003)
242. Manus ungual I, length: less than 25% the length of the radius (0); greater than 30% the length of the radius (1).
243. Manus unguals II and III, shape: transversely compressed and pointed (0); dorsoventrally compressed, with a rounded tip (1). (Modified from Norman, 2002; Weishampel et al., 2003)
244. Ilium, medial flange partly closes acetabulum: present (0); absent (1).
245. Ilium, acetabulum size; large and concave, extends deeply into body of ilium (0); very narrow (1). (Modified from Weishampel et al., 2003)
246. Ilium, relative lengths of pubic and ischial peduncles: extends ventrally to the same level (0); ischial peduncle extends beyond the pubic peduncle (1).
247. Ilium, Ischial peduncle extends distinctly laterally from the body of the ilium: absent (0); present (1).
248. o Ilium, ischial peduncle morphology: formed by a single large ventral protrusion (0); composed of a large oval ventral protrusion and by a smaller, caudodorsally located prominence (1); formed by two protrusions of similar size (2). (Modified from Prieto-Márquez and Salinas, 2010)
249. Ilium, preacetabular process, orientation; external surface faces laterally, in approximately the same plane as the iliac body (0); process is distinctly twisted about its long axis (1). (Weishampel et al., 2003)
250. Ilium, preacetabular process, lateral deflection relative to midline: 10-20 degrees (0); more than 30 degrees (1). (Butler et al., 2008)
251. Ilium, preacetabular process, morphology of distal end: parallel-sided or slightly tapering (0); small ventral flange (1).
252. Ilium, cranial sacral facets transversely widened so that the base of the preacetabular process is thicker ventrally than dorsally: absent (0); present (1).
253. Ilium, preacetabular process, ridge on medial side originating from medial sacral ridge: absent (0); present (1).
254. Ilium, preacetabular process, second medial ridge starting from the ventral edge of the process, along the notch above the pubic peduncle: absent, (0); present (1).
255. Ilium, dorsal margin of iliac body, shape in lateral view: straight or convex (0); concave (1).
256. Ilium, dorsal margin of the iliac body, shape in dorsal view: narrow, not transversely expanded (0); transversely expanded laterally to form a narrow shelf (1). (Modified from Butler et al., 2008)
257. Ilium, pendant process ("pendule" sensu Norman, 2015) extending from lateral ilial shelf dorsal to the ischial peduncle: absent (0); present (1).
258. Ilium, brevis shelf: present (0); absent (1).
259. Ilium, brevis fossa, well defined by a lateral lip: absent (0); present (1).
260. Ilium, brevis shelf and fossa, orientation: fossa faces ventrolaterally and shelf is near vertical, creating a deep postacetabular portion (0); fossa faces ventrally and shelf is horizontal (1). (Modified from Butler et al., 2008)
261. Ilium, ridge originating from the ischial peduncle and continuing caudally along the brevis shelf, often disappearing around the middle of the postacetabular process (this is distinct from the medial boundary of the brevis shelf): absent (0); present (1).
262. o Ilium, postacetabular process, width as a percentage of its length: less than 30% (0); between 35 and 55% (1); over 60% (2).
263. o Ilium, postacetabular process, length as a percentage of the total length of the ilium: 20% or less (0); 23-30% (1); 34% or more (2). (Modified from Butler et al., 2008)
264. Ilium, postacetabular process, shape: dorsal and ventral margins are parallel and roughly the same length, so the process has a square or rounded caudal end (0); the dorsal margin is shorter than the ventral, so that the caudal margin slopes caudoventrally (1); the dorsal and ventral edges slope smoothly towards each other, forming a pointed caudal end (2).
265. Ilium, postacetabular process, orientation: extends subhorizontally, does not extend above the body of the ilium (0); extends caudodorsally, so the postacetabular process reaches further dorsally than the body of the ilium (1).
266. Ilium, orientation of the medial sacral ridge: craniocaudally directed, parallel to the dorsal margin of the ilium, and ending caudal to the level of the ischial peduncle, well into the proximal region of the postacetabular process (0); diagonal, cranioventrally to caudodorsally oriented, concave ventrally and converging with the dorsal margin at the level of the ischial peduncle (1). (Prieto-Márquez and Salinas, 2010)
267. Pubis, prepubic process: absent (0); present (1). (Butler et al., 2008)
268. Pubis, prepubic process, width: compressed mediolaterally, dorsoventral height exceeds mediolateral width (0); mediolateral width exceeds dorsoventral height (1). (Butler et al., 2008)
269. Pubis, prepubic process, shape in lateral view: unexpanded distally (0); constricted proximal portion followed by a distal expansion (1). (Modified from Norman, 2002)
270. Pubis, prepubic process, depth of the dorsoventral expansion of the distal region greater than the width of the acetabular margin; absent (0); present (1). (Modified from Prieto-Márquez and Salinas, 2010)
271. o Pubis, prepubic process, craniocaudal length of the proximal constriction relative to the length of the dorsoventral expansion: constriction longer than the dorsoventral expansion (0); constriction and distal expansion subequal (1); constriction shorter than the dorsoventral expansion (2). (Modified from Horner et al., 2004; Prieto-Márquez and Salinas, 2010)
272. Pubis, prepubic process shape, if unexpanded distally: straight (0); concave dorsally (1).
273. Pubis, prepubic process, horizontal ridge on medial side: absent (0); present (1).
274. Pubis, prepubic process, extends beyond distal end of preacetabular process of ilium: absent (0); present (1). (Butler et al., 2008)
275. Pubis, obturator foramen completely enclosed: present (0); absent (1).
276. Pubis, obturator foramen shape: elliptical (0); circular (1).
277. Pubis, obturator foramen, orientation when the prepubis is oriented in a parasagittal plane: the obturator foramen faces laterally (0); the obturator foramen faces somewhat dorsally (1).
278. o Pubis, shaft (postpubis), length: approximately equal in length to the ischium (0); reduced, extends for about half the length of the ischium (1); very short, extends less than 20% the length of the ischium (2). (Modified from Butler et al., 2008)
279. Ischium, iliac peduncle, orientation of the acetabular and caudodorsal margins: divergent approaching the articulation with the ilium (0); either parallel or slightly convergent (1). (Prieto-Márquez and Salinas, 2010)
280. Ischium, pubic peduncle, shape: transversely compressed (0); dorsoventrally compressed (1). (Butler et al., 2008)
281. Ischium, shaft, shape in lateral view: straight (0); downwardly curved (1). (Butler et al., 2008; Weishampel et al., 2003; Norman, 2002)
282. Ischium, shaft, orientation: axis of shaft is caudoventral (0); axis of shaft is caudal (1).
283. Ischium, shaft, twisted along its length so that the lateral side faces ventrally on the distal part of the shaft: present (0); absent (1).
284. Ischium, cross-section at mid-shaft: compressed mediolaterally (0); ovoid or subcircular (1). (Modified from Weishampel et al., 2002, and Butler et al., 2008)
285. Ischium, obturator process: absent (0); present (1). (Modified from Weishampel et al., 2003 and Butler et al., 2008)
286. Ischium, obturator process, shape: discrete tab (0); elongate ridge extending distally down ischial shaft (1).
287. Ischium, obturator process, position of proximal edge: beyond the proximal 29% of the length of the ischium (measured from the acetabular rim to the distal end) (0); within the proximal 25% (1). (Modified from Weishampel et al., 2003)
288. Ischium, distal end expanded to 50% or more the depth of the adjacent ischial shaft; absent (0); present (1). (Modified from Weishampel et al., 2003, Norman, 2002)
289. *Femur, length relative to tibia: shorter than or equal to (0); longer than the tibia (1). (Milner and Norman, 1984; Norman, 1984b, modified from Weishampel et al., 2003)
290. *Femur, shape in lateral view: straight (0); bowed cranially (1). (Butler et al., 2008)
291. Femur, shape in cranial view: straight (0); bowed laterally (1). (Weishampel et al., 2003)
292. Femur, head: confluent with greater trochanter (0); distinct constriction separates head and greater trochanter (1). (Butler et al., 2008)
293. Femur, head, ventromedially oriented sulcus on caudal side: absent (0); present (1).
294. Femur, 'anterior' or 'lesser' trochanter, craniocaudal width: subequal to that of the greater trochanter (0); substantially less than that of the greater trochanter (1). (Modified from Butler et al., 2008)
295. Femur, anterior trochanter, level of most proximal point relative to level of proximal femoral head: positioned distally on the shaft (0); positioned proximally, approaches level of proximal surface of femoral head (1). (Modified from Butler et al., 2008)
296. Femur, fourth trochanter, shape: pendant (0); prominent ridge (1). (Butler et al., 2008; Weishampel et al., 2003; Norman, 2002)
297. Femur, fourth trochanter, if ridge-shaped, lateral profile of the caudoventral margin: triangular and ending in a caudally, and slightly ventrally, directed point (0); smooth and arcuate (1). (Modified from Prieto-Márquez and Salinas, 2010)
298. *Femur, fourth trochanter, position: located entirely on proximal half of femur (0); positioned at midlength or distal to midlength (1). (Weishampel et al., 2003; Butler et al., 2008)
299. Femur, muscle scar for M. caudofemoralis longus on medial side of fourth trochanter, position and size: a small, distinct fossa on the shaft of the femur (0); a large oblong depression mainly on the medial side of the fourth trochanter (1).
300. Femur, cranial (extensor) intercondylar sulcus on distal end: absent (0); present (1). (Norman, 2002; Weishampel et al., 2003; Butler et al., 2008)
301. o Femur, cranial intercondylar sulcus, shape: open, U-shaped (0); partially enclosed by cranial expansion of condyles (1); fully enclosed by cranial condyles (2). (Norman, 2002)
302. *Femur, caudal (flexor) intercondylar sulcus of the femur: fully open (0); medial condyle inflated laterally, partially covers opening of flexor sulcus (1). (Weishampel et al., 2003; Butler et al., 2008).
303. Femur, lateral (fibular) condyle, position and size: not inset from the lateral edge, and only slightly narrower in width than the medial condyle (0); strongly inset medially, reduced in width relative to medial condyle (1). (Modified from Butler, 2008)
304. Femur, distal condyles, shape in lateral view: moderately expanded craniocaudally (0); strongly expanded (condyle extends cranially as well as caudally) (1). (Modified from Norman, 2002)
305. Femur, distal condyles, shape of articular surface: flat articular surface (0); rounded articular surface (1); flat to concave lateral condyle and convex medial condyle (2).
306. Tibia, fibular process shape: single (0); double (1).
307. Tibia, expansion of proximal end in lateral view: width of proximal tibia is less than 2.3 times the diameter at midshaft (0); proximal tibia is more than 2.5 times the diameter at midshaft (1).
308. Fibula, expansion of proximal end, shape: flared, with concave cranial and caudal margins (0); nearly straight cranial and caudal margins (1).
309. Astragalus, astragular notch on lateral margin of ascending process: absent (0); present (1).
310. Astragalus, small fossa on cranial side of astragalus (this is directly ventral to astragalar notch when both are present): absent (0); present (1).
311. Astragalus, joint with calcaneum, morphology: peg-in-socket (0); straight butt joint (1).
312. Astragalus, width relative to calcaneum width: less than 2.25 times calcaneum width (0); greater than 2.5 times calcaneum width (1).
313. Distal tarsals II and III: present (0); absent (1). (Horner et al., 2004, character 102; Prieto-Márquez and Salinas, 2010)
314. Medial distal tarsal: articulates distally with metatarsal III only (0); articulates distally with metatarsals II and III (1). (Butler et al., 2008)
315. Metatarsal I: absent (0); present (1).
316. Metatarsal I, bears digits: present (0); absent (1). (Norman, 2002; Weishampel et al., 2003; Butler et al., 2008)
317. Digit I, length: metatarsal I robust and well-developed, distal end of phalanx I-1 projects beyond the distal end of metatarsal II (0); metatarsal I reduced and proximally splint-like, end of phalanx I-1 does not extend beyond the end of metatarsal II (1). (Modified from Butler 2008)
318. Metatarsal II, tab-like process on craniolateral edge that articulates with metatarsal III: absent (0); present (1).
319. Metatarsal III, ratio of length to mediolateral width at midshaft: less than 6.5 (0); greater than 7.5 (1). (Modified from Prieto-Márquez and Salinas, 2010)
320. Metatarsal V: present (0); absent (1). (Modified from Weishampel et al., 2003; Milner and Norman, 1984; Sereno, 1984; Norman, 1986; Coria and Salgado, 1996)
321. o Pedal unguals, shape: mediolaterally compressed, tapering, and pointed (0); dorsoventrally compressed, proximally wider, tapering to a bluntly truncated tip (1); dorsoventrally compressed, mediolaterally broad and proximodistally shortened, rounded distal end ("hoof-shaped") (2). (Modified from Prieto-Márquez and Salinas, 2010; Butler et al., 2008; Weishampel et al., 2003; Norman, 2002)
322. Tendons, ossified hypaxial tendons on caudal vertebrae: absent (0); present (1). (Butler et al., 2008)
323. Tendons, ossified epaxial and hypaxial tendon arrangement: longitudinal (0); double-layered lattice (1). (Butler et al., 2008).
APPENDIX 2.
Character matrix (also available as TNT and Nexus files in supplementary documents) for download.
APPENDIX 3
List of taxa used in analysis and sources from which characters were scored.
Institutional Abbreviations: AM, Albany Museum, Grahamstown, South Africa; AMNH, American Museum of Natural History, New York City, New York, USA; BYU, Earth Sciences Museum, Brigham Young University, Provo, Utah, USA; CEUM, College of Eastern Utah Prehistoric Museum, Price, Utah, USA; CM, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA; DMNS, Denver Museum of Nature and Science, Denver, Colorado, USA; GPIT, Institut und Museum für Geologie und Paläontologie der Universität Tübingen, Tübingen, Germany; IVPP, Institute of Vertebrate Paleontology and Paleoanthropology, Beijing, China; MB, Museum für Naturkunde Berlin, Berlin, Germany; MC, Museum Crúzy, Crúzy, France; MCF, Museo Carmen Funes, Plaza Huincul, Neuquén, Argentina; MCS, Museo de Cinco Saltos, Rio Negro Province, Argentina; MHN-AIX-PV, Museum d’Histoire Naturelle d’Aix-en-Provence, Aix-en-Provence, France; MHNM, Museum d’Histoire Naturelle de Marseille, Marseille, France; MNHN, Museum National d’Histoire Naturelle, Paris, France; MOR, Museum of the Rockies, Bozeman, Montana, USA; MUCPv, Museo de Geologia y Paleontologia de la Universidad Nacional del Comahue, Neuquén, Argentina; NHMUK, Natural History Museum, London, United Kingdom; NMV, National Museum of Victoria, Melbourne, Australia; OUMNH, Oxford University Museum of Natural History, Oxford, UK; QM, Queensland Museum, Geoscience Collection, Brisbane, Queensland, Australia; RBINS (formerly IRSNB), Royal Belgian Institute of Natural Sciences, Brussels, Belgium; ROM, Royal Ontario Museum, Toronto, Canada; SAM, South African Museum (Iziko Museums of Cape Town), Cape Town, South Africa; SDSM, Museum of Geology, South Dakota School of Mines and Technology, Rapid City, South Dakota, USA; UMNH, Natural History Museum of Utah, Salt Lake City, Utah, USA; USNM, United States National Museum of Natural History, Washington, DC, USA; YPM, Yale Peabody Museum, New Haven, Connecticut, USA; ZDM, Zigong Dinosaur Museum, Dashanpu, China.
| Taxon | Specimen Numbers | Literature Sources |
|---|---|---|
| Agilisaurus | Photographs of holotype specimens ZDM T6011 kindly provided by Han Fenglu. | Barrett et al., 2005 |
| Altirhinus kurzanovi | Norman, 2002 | |
| Anabisetia saldiviai | MCF-PVPH-74-6. | Coria and Calvo, 2002 |
| Atlascopcosaurus | NMV P166409 (holotype), 182967. | |
| Bactrosaurus | Weishampel and Horner, 1986; Godefroit et al., 1998 | |
| Barilium | NHMUK R798a R798b, R799-806, R4742, 4771: All holotype specimen. | Norman, 2010 |
| Batyrosaurus | Godefroit et al., 2012 | |
| Bolong | Wu and Godefroit, 2012 | |
| Camptosaurus | YPM 1877 (holotype), 1878, USNM V4277, 5473, UMNH.VP.5322, 16422, 16452, 16454, 16455, 16458, 16572, 16592-3. | Gilmore, 1909; Brill and Carpenter, 2006 |
| Cedrorestes | DMNS 47994 (holotype). | |
| Cumnoria | OUMNH Geol J3303 (holotype). | |
| Dakotadon | SDSM 8651 (holotype). | |
| Dryosaurus | CM 3392, 1949, 87688, DMNS 9001 (juvenile). | Galton, 1981, 1983 |
| Dysalotosaurus | MB R1316, 1318, 1320, 1322, 1329, 1333, 1335, 1338, 1351, 1358, 1370, 1372-8, 1383, 1394, 1396-8, 1408-9, 1474, 1476, 1478, 1480-1, 1485-7, 1502, 1540, 1565-6, 1585-1604, 1707, 1709, 1711, 1717-8, 2144, 2503, 2508, 2511, 2516-7, 3293, 3297, 3468, 3474. GPIT/RE/3608, 3612, 3614, 3826, 3829, 3830, 4156, 4181, 4189, 4545, 4710, 4925, 5192, 5302, 5355, 5460-3, 5636, 6067, 6161, 6169, 6189, 6269, 6399, 6544, 6549, 6803, 6987, 8425. | Janensch, 1955 |
| Edmontosaurus | Lambe, 1920; Campione and Evans, 2011 | |
| Eocursor | SAM PK K8025 (holotype). | Butler et al., 2007 |
| Eolambia | CEUM 9758 (holotype), 1233, 1251-1256, 3207, 3186, 13383, 14419, 14464, 14466, 14495, 14534, 14576, 14601, 14602, 34247, 34252, 34329, 4337, 34382, 34397, 34412, 34430, 34441, 35323, 35384, 35412, 35413, 35492, 35524, 35525, 35539, 35592, 35637, 35638, 35673, 35679, 35733, 35742, 36190, 52097, 52865, 52884, 52924, 52936, 52988, 73585, 74566, 74572, 74611, 74653, 78380. | McDonald et al., 2012 |
| Equijubus | IVPP V 12534 | You et al., 2003 |
| Fukuisaurus | Kobayashi and Azuma, 2003 | |
| Gasparinisaura | MCS 1-3; MUCPv 111-2, 208 (holotype), 210, 212-3 | Coria and Salgado, 1996 |
| Gilmoreosaurus | Weishampel and Horner, 1986; Prieto-Márquez and Norrell, 2010 | |
| Shishugou taxon | IVPP 14559, field number field number WCW-02-29 | |
| Haya | Makovicky et al., 2011 | |
| Hexinlusaurus (=Yandusaurus) multidens | Photos of holotype ZDM T6001 kindly provided by Han Fenglu. | Barrett et al., 2005 |
| Hippodraco | UMNH VP 20208 (holotype) | McDonald et al., 2010b |
| Hypacrosaurus | MOR 549 (holotype H. stebingeri), MOR 355, MOR 548. | |
| Hypselospinus | NHMUK R1650 (holotype), 33, 1627, 1636, 1831, 1834. Iguanodon hollingtoniensis holotype: NHMUK R604, 811, 811b, 1148, 1629, 1632. | Norman, 2010, 2014 |
| Hypsilophodon | NHMUK R146, 192-4, 197 (holotype), 2466, 2477, 2488. | Galton, 1974 |
| Iguanacolossus | UMNH VP 20205 (holotype) | McDonald et al., 2010b |
| Iguanodon | RBINS R51 (lectotype) | Norman, 1980 |
| Jeholosaurus | Barrett and Han, 2009; Han et al., 2012 | |
| Jeyawati | McDonald et al., 2010a | |
| Jinzhousaurus | IVPP V12691 (holotype) | Barrett et al., 2009; Wang et al., 2010 |
| Kangnasaurus | SAM 2732 (holotype), 2731a-j | Cooper, 1985 |
| Kirkwood taxon | AM 6150 (holotype), 6004, 6005, 6021, 6022, 6030, 6053, 6055, 6056, 6063, 6066, 6067, 6077, 6093, 6101, 6102, 6103, 6104, 6107, 6108, 6109, 6110, 6111, 6119, 6122, 6151, 6154, 6155, 6175, 6176, 6190 | |
| Kukufeldia | NHMUK R28660 | |
| Lanzhousaurus | You et al., 2005 | |
| Leaellynasaura | NMV P185990-1 (holotype), 185992-3, 186047, 221080, 229196 | |
| Lesothosaurus | NHMUK R8501, NHMUK RUB23, SAM PK K 400, 401, 1106 | Sereno, 1991; Butler, 2005 |
| Levnesovia | Sues and Averianov, 2009 | |
| Lurdusaurus | MNHN.F.GDF 1700 (holotype) | |
| Macrogryphosaurus | MUCPv 321 (holotype) | Calvo et al., 2007 |
| Maiasaura | Horner 1983; Dilkes, 2000 | |
| Mantellisaurus | NHMUK R5764 (holotype); RBINS R57 (this specimen was scored as a separate OTU). | Norman, 1986 |
| Muttaburrasaurus | QM F6140 (holotype), 14921 | |
| Orodromeus | MOR 294 (holotype); MOR 1141; MOR 473 | |
| Oryctodromeus | MOR 1636 | Varricchio et al., 2007 |
| Othnielosaurus | BYU-ESM-163 | |
| Ouranosaurus | MNHN.F.GDF 300 (casts) | |
| Owenodon | NHMUK R2998 | |
| Parksosaurus | Photographs of ROM 804 (holotype) provided by Ali Nabavizadeh | Parks, 1926 |
| Planicoxa | DMNS 42504 (holotype), 40909, 40914, 40917, 40918, 42477, 42505, 42506, 42507, 42508, 42511, 42513, 42521, 42525, 42599. | |
| Proa | McDonald et al., 2012 | |
| Probactrosaurus | Norman, 2002 | |
| Protohadros | Head, 1998; Main, 2013 | |
| Rhabdodon | MHNM.6034.11998 (holotype), MHNAIX-PV.1995, MHN-AIX-PV.2007, MCM30, 1252, 1575, 2111 3036, 3583, 4058, MC-MN30, 32, 36, 365, MC-MOB6, 7013, MC-PSP3, MC-CY.QR.1-6, 8, 11, 15, 18, 21, 24 | Chanthasit, 2010 |
| Shuangmiaosaurus | You et al., 2003 | |
| Stormbergia | SAM PK K1105 (holotype), 1107 | Butler, 2005 |
| Tenontosaurus dossi | Winkler et al., 1997 | |
| Tenontosaurus tilletti | YPM 5456, 5459, AMNH 3031, 3034, 3040 | |
| Talenkauen | Novas et al., 2004; Cambiaso, 2007 | |
| Telmatosaurus | NHMUK R3386, 3388, 4911 | Weishampel et al., 1993 |
| Tethyshadros | Dalla Vecchia, 2010 | |
| Theiophytalia | YPM 1887 | Brill and Carpenter, 2006 |
| Thescelosaurus infernalis | SDSM 7210 (holotype): MOR 979 | |
| Thescelosaurus neglectus | AMNH 117, 5030-1, 5034 | Boyd et al., 2009; Boyd 2012 |
| Trinisaura | Coria et al., 2013 | |
| Uteodon | CM 11337 (holotype), 15780 (paratype), 21723, 79050 (paratype) | Carpenter and Wilson, 2008; McDonald, 2011 |
| Valdosaurus | Barrett et al., 2011 | |
| Xuwulong | You et al., 2011 | |
| Zalmoxes robustus | NHMUK R3809-10, 3812-4, 3390, 3392 (holotype), 3393, 3836, 4912 | Weishampel et al., 2003 |
| Zalmoxes shqiperorum | NHMUK R4900 (holotype) | Weishampel et al., 2003 |
| Zephyrosaurus | Sues, 1980 |
APPENDIX 4
The following taxa are excluded from the analysis. In some cases, these are junior synonyms of taxa that are included, while other taxa lack apomorphies or a distinct combination of synapomorphies.
Darwinsaurus evolutionis Paul, 2012
= Hypselospinus fittoni Norman, 2010
= Iguanodon fittoni Lydekker, 1889
Type horizon, locality, and age: Wadhurst Clay Formation, England, Valanginian
Holotype: NHMUK R1831, 1833, 1835, 1836
This study follows Norman (2013, 2015) in considering this species a nomen dubium, based on the incorrect anatomical details (e.g., a long diastema) described in the dentary of NHMUK R1831, and the disparate localities of the designated holotype. Specimens NHMUK R1831, 1833, and 1835 from the Valanginian Wadhurst Clay Formation are referred to Hypselospinus fittoni, while NHMUK R1836 from the Barremian of the Isle of Wight is referred to Mantellisaurus atherfieldensis (Norman, 2014b). There has been much dispute about the presence of a diastema in this specimen (Norman, 2010; Paul, 2012; Norman, 2014b); firsthand examination by the author finds that the area rostral to the preserved teeth is weathered, but alveoli are clearly present. This highlights the importance of examining specimens directly. No matter how detailed a drawing or photograph may appear to be, they do not always reproduce the necessary details of a specimen.
Dollodon bampingi Paul, 2008
= Mantellisaurus atherfieldensis (Hooley, 1925)
Type horizon, locality, and age: Upper Hainaut Group, Belgium, late Barremian to early Aptian
Holotype: RBINS R57
The holotype of this species had previously been referred to Mantellisaurus (= Iguanodon) atherfieldensis. McDonald (2012b) and Norman (2013, 2015), both consider it to be a nomen dubium, though if the taxon is not valid it should be considered a junior subjective synonym of M. atherfieldensis. In order to further test whether this specimen and the holotype of M. atherfieldensis form a monophyletic group, they were coded as separate OTUs in this analysis.
Delapparentia turolensis Ruiz-Omenaca, 2011
= Iguanodon bernissartensis Boulanger, 1881
Type horizon, locality, and age: Camarillas Formation, Teruel, Spain, early Barremian
Holotype: MPT/I.G.
Norman (2014b) considered this taxon a nomen dubium, however Gasca et al. (2015) offered an emended diagnosis based on new material. These new characters, however, fail to distinguish Delapparentia from Barilium. Gasca et al. (2015) cite a single autapomorphy, a neural spine of the axis with a height from the base of the postzygapophysis that is greater than half the length of the neural spine. However, the neural spine is clearly broken cranially, so the length in this specimen cannot be determined. The unique combination of characters listed by Gasca et al. (2015) are all present in Barilium: the preacetabular process twists along its length so the lateral surface faces dorsally at the distal end, a large process is present and visible in the preacetabular notch of the ilium for articulation with the sacral rib, a straight dorsal margin of the ilium, a distally expanded prepubic process, and proximal caudal vertebral centra dorsoventrally expanded. While it is clear that an iguanodontian similar to or synonymous with Barilium dawsoni was present in the Barremian of Spain, the species D. turolensis remains inadequately diagnosed, and is not included in this analysis.
Elrhazosaurus (=Valdosaurus) nigerensis (Galton and Taquet, 1982)
Type horizon, locality, and age: Elrhaz Formation, Niger, late Aptian
Holotype: MNHN GDF 332
This species, assigned to its own genus by Galton (2009), is erected on a single femur. Galton (2009) assigns it to Dryosauridae based on the deep pit on the medial side of the femur, just cranial to the fourth trochanter. However, this character state is found widely among basal neornithischians, in genera such as Anabisetia, Hexinlusaurus, and Orodromeus. The other features cited by Galton (2009) in the diagnosis of the new genus are plesiomorphic within Ornithopoda (greater trochanter extends further dorsally than lesser trochanter) or vague (“transversely wide raised area separating deep pit from base of fourth trochanter”), except for the deep and obliquely inclined extensor groove. This feature is similar to the extensor grooves in Valdosaurus and Dysalotosaurus, and therefore not sufficient to diagnose the species. Elrhazosaurus nigerensis is here considered a nomen dubium.
Huxleysaurus (=Iguanodon) hollingtoniensis (Lydekker, 1889)
= Hypselospinus fittoni, Norman, 2010
Type horizon, locality, and age: Wadhurst Clay Formation, England, Valanginian
Holotype: NHUMK R1148, 1629, 1632, 811, 811b, 604.
The genus Huxleysaurus was named by Paul (2012) based on the holotype specimen of Iguanodon hollingtoniensis, which was previously referred to Hypselospinus fittoni (Norman, 2010). While there is fairly little overlapping material between this specimen and that of the holotype of Hypselospinus fittoni, the areas of overlap are consistent with one another, including a longitudinal ridge along the brevis shelf (Character 261). Furthermore, the diagnosis of Hu. hollingtoniensis is inaccurate and entirely inadequate. In full, it reads “Femur robust, moderately curved, 4th trochanter pendent.” While the fourth trochanter of this specimen does come to a sharp point at its caudoventral margin, it is not pendant. While Norman (2014b) considers Hu. hollingtonensis a nomen dubium, it is here considered a junior subjective synonym of Hypselospinus fittoni.
Mantellodon carpenteri Paul, 2012
= Mantellisaurus atherfieldensis (Hooley, 1925)
Type horizon, locality, and age: Lower Greensand Formation, England, early Aptian
Holotype: NHMUK R3791
This new genus and species was described based on the so-called Maidstone specimen, now on display at the Natural History Museum in London. The diagnosis of this taxon, in full, states: “Limb elements slender. Ilium deep, anterior process robust, posterior acetabular body short and very triangular, dorsal margin strongly arched.” This diagnosis is vague and does not differentiate this specimen from others assigned to Mantellisaurus atherfieldensis, or even from other genera such as Ouranosaurus. It is here considered a junior subjective synonym of M. atherfieldensis.
Kukufeldia tilgatensis McDonald, Barrett, and Chapman, 2010
Type horizon, locality, and age: Tunbridge Wells Sand Formation, Wealden Group, England, middle to late Valanginian
Holotype: NHMUK R28660
This taxon, based on the single dentary NHMUK R28860, was diagnosed based on the autapomorphy of a distinct row of foramina. While it is recognized as a nomen dubium by Norman (2014b), who also cites that McDonald now agrees with this, it remains unclear whether this specimen can be assigned to Barilium or to Iguanodon. As such, it was coded as a distinct OTU in this analysis.
Osmakasaurus depressus (Gilmore, 1909)
= Camptosaurus depressus Gilmore, 1909
= Planicoxa depressa Carpenter and Wilson, 2008
Type horizon, locality, and age: Lakota Formation, South Dakota, Barremian-Aptian
Holotype: USNM 4753
This genus was erected by McDonald (2011). The specific diagnosis for this taxon is by a unique combination of characters: distal end of preacetabular process of the ilium with a ventral flange (“horizontal boot” of McDonald, 2011), straight dorsal margin of the ilium, and the dorsal margin of the ilium transversely expanded. However, these character states appear widely in Iguanodontia, and are found together in several genera, including Mantellisaurus and Iguanacolossus. Thus, Osmakasaurus depressus is considered a nomen dubium.
Penelopognathus weishampeli Godefroit, Li and Shang, 2005
Type horizon, locality, and age: Bayan Gobi Formation, Inner Mongolia, China, Albian
Holotype: IMM 2002-BYGB-1
This species is erected on a single dentary. As it displays only characters that are widespread among styracosternans, it is here considered a nomen dubium, and not used in the analysis. This is supported by Norman (2014b), who noted that the diagnosis contains no unique characters.
Proplanicoxa galtoni Carpenter and Ishida, 2010
Type horizon, locality, and age: Upper Wessex Formation, England, late Barremian
Holotype: NHMUK R8649
This species is diagnosed by a postacetabular process of the ilium that is directed about 50° dorsolaterally. It is clear based on the wide variety of shapes present among the ilia in the specimens of Iguanodon bernissartensis that these elements are prone to post-mortem distortion, and this single character state is insufficient to diagnose a species. It is here considered a nomen dubium.
Qantassaurus intrepidus Rich and Rich, 1999
Type horizon, locality, and age: Wonthaggi Formation, Victoria, Australia, early Aptian
Holotype: NMV P199075
This taxon is diagnosed based on the deep, foreshortened dentary with only 10 alveoli. However, a low tooth count is found in juvenile individuals among many neornithischians (Horner and Currie, 1994; Scheetz, 1999; Hübner and Rauhut, 2010), and the dorsoventrally deep dentary appears to be a taphonomic feature caused by mediolateral crushing of a transversely wide dentary. The individuals referred to this taxon most likely pertain to the coeval Atlascopcosaurus loadsi, though the fragmentary nature of the specimens makes this difficult to determine.
Sellacoxa pauli Carpenter and Ishida, 2010
Type horizon, locality, and age: Lower Wadhurst Clay, England, early Valanginian
Holotype: NHMUK R3788
The holotype of this species has previously been referred to Iguanodon dawsoni, and later to Barilium dawsoni (Norman 2010, 2015). While this specimen lacks some features used to diagnose B. dawsoni, such as the transversely wide articular facet for the sacrum visible laterally through the preacetabular notch, it is also not clear that it is a distinct taxon. The diagnosis of Carpenter and Ishida includes one character state that cannot be properly determined: the sharply cranioventrally sloped preacetabular process. This region is reconstructed with plaster near its base, so the actual angle is unclear. The other characters cited are all found widely within ankylopollexians (dorsal margin of ilium concave, body of ilium deep, ischial peduncle extends laterally onto body of ilium, prepubic process mediolaterally compressed and expanded only at the distal end, ischium “T-shaped” with a straight shaft). While the proper taxonomic position of NHMUK R3788 remains unclear, the name Sellacoxa pauli is here considered a nomen dubium.
Ratchasimasaurus suranareae Shibata, Jintaskul and Azuma, 2011
Type horizon, locality, and age: Khok Kraut Formation, Thailand, Aptian
Holotype: NRRU-A2064
Siamodon nimngami Buffetaut and Suteethorn, 2011
Type horizon, locality, and age: Khok Kraut Formation, Thailand, Aptian
Holotype: PRC-4
Both of these species were named based on non-overlapping material from the Aptian-aged Khok Kraut Formation of Thailand. While they could offer intriguing biogeographic information, the fragmentary specimens reveal only characters that are widespread within Styracosterna, and are not included in the current analysis.
APPENDIX 5
Unambiguous synapomorphies of major clades. Where topology differs between parsimony and Bayesian analyses, these are listed separately.
| Taxon | Unambiguous synapomorphies (Character numbers and states correspond to those given in Appendix 1.) |
| Thescelosauridae (Parsimony) | 98.0 Dentary, shape of tooth row in dorsal view: straight.
136.3 Maxillary teeth, maximum number of ridges extending from the base to the tip of the crown on labial side of teeth: ten to twelve ridges. 196.0 Scapula, acromion process length: does not extend beyond the edge of the coracoid. |
| Thescelosauridae (Bayesian) | 136.3 Maxillary teeth, maximum number of ridges extending from the base to the tip of the crown on labial side of teeth: ten to twelve.
137.3 Dentary teeth, maximum number of ridges extending from the base to the tip of the crown on lingual side of teeth: nine to eleven. |
| Clypeodonta (Parsimony) | 63.1 Quadrate buttress ("hamular process"): present.
69.2 Quadrate, mandibular condyle proportions: lateral condyle is larger than medial. 70.0/1 Quadrate, mandibular condyles, orientation of distal margin relative to long axis of quadrate: dorsomedially sloped or horizontal. 127.1 Maxillary teeth, morphology of roots: crown tapers to root. 128.1 Dentary teeth, morphology of roots: crown tapers to root. 139.1 Maxillary teeth, primary ridge on labial side of crown: present. 146.1 Maxillary teeth, base of crown defined by an everted lip which makes the crown slightly inset from the root: present. 147.1 Dentary teeth, base of crown defined by an everted lip which makes the crown slightly inset from the root: present. 159.1 Postaxial cervical vertebrae, proportions of centra: elongate, with craniocaudal length more than twice the dorsoventral height. |
| Clypeodonta (Bayesian) | 16.1 Maxilla, diastema: present, first maxillary tooth is at least one crown width from the premaxilla.
84.1 Predentary, length of oral margin: subequal in length to the oral margin of the premaxilla. 101.1 Dentary, coronoid process, height: extends more than one crown height above tooth row. 111.1 Surangular, small fenestra positioned dorsally on or near the dentary joint: present. 114.1 Surangular foramen, position relative to the lateral lip of the glenoid: rostral. 134.1 Cheek teeth, enamel distribution: asymmetrical: thicker on the labial side of maxillary teeth and on the lingual side of dentary teeth. 135.1 Cheek teeth, apicobasally extending ridges on labial side of maxillary teeth and lingual side of dentary teeth: present. 292.1 Femur, head: distinct constriction separates head and greater trochanter. |
| Iguanodontia (Parsimony) | 21.1 Maxilla, dorsal process, shape in lateral view: broad and triangular.
60.1 Quadratojugal, position of ventral margin relative to mandibular condyle of the quadrate: well-removed from the mandibular condyle. *131.1 Cheek teeth: number of wear facets on each tooth: one. 157.1 Postaxial cervical vertebrae, form of central surfaces: at least slightly opisthocoelous. 185.1 Caudal vertebrae, length of prezygopophyses on distal caudals: elongate, extending nearly to the midpoint of the preceding vertebra. 188.1 Chevrons, shape: strongly asymmetrically expanded distally, width greater than length in mid caudals. 199.1 Scapula, supraglenoid fossa: present. *236.1 Manus digit III, number of phalanges: three or fewer. 263.1 Ilium, postacetabular process, length as a percentage of the total length of the ilium: 23-30%. |
| Iguanodontia (Bayesian) | 3.1 Premaxilla, shape of subnarial region: ventral premaxilla flares laterally to form a partial floor of the narial fossa.
31.1 Antorbital fenestra, size: relatively small (10% basal skull length or less). 75.1 Posttemporal foramen/fossa, position: forms a notch in the dorsal margin of the paroccipital process, enclosed dorsally by the squamosal. 87.1 Predentary, denticulate oral margin: present. 89.1 Predentary, ventral process, depth of bifurcation: deep bifurcation. 125.0 Cheek teeth; dentary teeth extend farther rostrally than maxillary teeth: absent. 126.1 Cheek teeth, spaces between the roots: absent, teeth are closely packed. 127.1 Maxillary teeth, morphology of roots: crown tapers to root. 128.1 Dentary teeth, morphology of roots: crown tapers to root. 129.1 Maxillary teeth, ratio of crown height to width (for unworn teeth): 1.25 to 1.9. *131.1 Cheek teeth: number of wear facets on each tooth: one. 148.1 Cheek teeth, at least moderately developed labiolingual expansion at the base of the crown ("cingulum"): absent. 177.0 Caudal vertebrae, position of the most distal caudal rib (transverse process): 12th caudal vertebrae or more proximal. 183.0 Caudal vertebrae, proportions of articular facets for chevrons: the distal facet on a given vertebra is much larger than the proximal facet. 214.1 Humerus, deltopectoral crest length: 43% of total humeral length or more. *236.1 Manus digit III, number of phalanges: three or fewer. 239.1 Phalanges of manual digits II-IV, length: first phalanx more than twice the length of the second phalanx. 283.1 Ischium, shaft, twisted along its length so that the lateral side faces ventrally on the distal part of the shaft: absent. 288.1 Ischium, distal end expanded to 50% or more the depth of the adjacent ischial shaft: present. 300.1 Femur, cranial (extensor) intercondylar sulcus on distal end: present. 302.1 Femur, caudal (flexor) intercondylar sulcus of the femur: medial condyle inflated laterally, partially covers opening of flexor sulcus (1). |
| Rhabdodontoidea(Parsimony) | 34.1 Orbit shape: subrectangular at least in its lower margin.
56.1 Jugal, shape of ventral edge: sinuous. 265.1 Ilium, postacetabular process orientation: extends caudodorsally, so the postacetabular process reaches further dorsally than the body of the ilium. 290.0 Femur, shape in lateral view: straight. |
| Rhabdodontoidea (Bayesian) | 21.1 Maxilla, dorsal process, shape in lateral view: broad and triangular.
26.2 Maxilla, shape of tooth row in ventral view: straight. |
| Tenontosaurus+ Rhabdodontidae (Parsimony) | 49.1 Jugal, type of suture with maxilla: 'finger-in-recess' joint.
55.1 Jugal, form of articulation with quadratojugal: tongue and-groove contact whereby the jugal lies lateral to the quadratojugal dorsally while ventrally the jugal lies medial to the quadratojugal. 65.0 Quadrate (paraquadratic) foramen or notch on boundary between quadrate and quadratojugal: absent. 123.0 Maxillary teeth, number; 13 or fewer tooth positions. 212.0 Humerus, deltopectoral crest form: a well-developed projection. 275.0 Pubis, obturator foramen completely enclosed: present. |
| Rhabdodontidae | 138.1 Cheek teeth: apicobasally extending ridges on cutting surface of unworn teeth (lingual surface of maxillary teeth, labial surface of dentary teeth): present.
196.0 Scapula, acromion process, length: does not extend beyond the edge of the coracoid. 216.2 Humerus, relative width of medial and lateral distal condyles: medial condyle wider. 219.1 Ulna, flange on proximal end that wraps around the lateral edge of the radius: present. 249.1 Ilium, preacetabular process, orientation: process is distinctly twisted about its long axis. 280.1 Ischium, pubic peduncle, shape: dorsoventrally compressed. 304.1 Femur, distal condyles, shape in lateral view: strongly expanded (condyle extends cranially as well as caudally). 320.1 Metatarsal V: absent. |
| Dryomorpha (Parsimony) | 6.1 Premaxilla, posterolateral process contacts lacrimal: present.
18.1 Maxilla, rostrolateral process in addition to the premaxillary process: present. 26.0 Maxilla, shape of tooth row in ventral view: medially bowed, with rostral and caudal ends curving laterally. 27.1 Maxilla, articulation with lacrimal: lacrimal fits into slot between medial and lateral portions of the dorsal process of the maxilla. 47.0 Jugal, exclusion of the rostral process from the antorbital fenestra by lacrimal-maxilla contact: absent. 57.1 Quadratojugal, shape: no dorsal process, element is small and blocky. 58.0 Quadratojugal, foramen through center of element: absent. 109.1 Coronoid process, relative craniocaudal widths of dentary and surangular (at midpoint of coronoid process): subequal. 145.1 Cheek teeth, secondary ridges, shape in cross-section: thin, sharp ridges, formed mainly by enamel. 224.1 Radius, shape in distal view: triangular. 287.1 Ischium, obturator process, position of proximal edge: within the proximal 25%. 322.0 Tendons, ossified hypaxial tendons on caudal vertebrae: absent. |
| Dryomorpha (Bayesian) | *6.1 Premaxilla, posterolateral process contacts lacrimal: present.
*18.1 Maxilla, rostrolateral process in addition to the premaxillary process: present. *27.1 Maxilla, articulation with lacrimal: lacrimal fits into slot between medial and lateral portions of the dorsal process of the maxilla. *57.1 Quadratojugal, shape: no dorsal process, element is small and blocky. 63.1 Quadrate buttress ("hamular process"): present. 65.1 Quadrate (paraquadratic) foramen or notch on boundary between quadrate and quadratojugal: present. 90.1 Predentary, short midline process dorsal to dentary symphysis: present. *109.1 Coronoid process, relative craniocaudal widths of dentary and surangular (at midpoint of coronoid process): subequal. *145.1 Cheek teeth, secondary ridges, shape in cross-section: thin, sharp ridges, formed mainly by enamel. *224.1 Radius, shape in distal view: triangular. *287.1 Ischium, obturator process, position of proximal edge: within the proximal 25%. |
| Dryosauridae | 9.1 Premaxilla, medial dorsal (nasal) process contacts the nasal: absent.
42.1 Palpebral(s), length relative to rostrocaudal width of orbit if unfused to orbital rim: traverses entire width of orbit. 97.0 Dentary, relationship of dorsal and ventral margins (under the tooth row): converge anteriorly. 143.0 Maxillary teeth, primary ridge position: centered, although some teeth within the same dental battery may display a slight offset of the primary ridge. 161.0 Dorsal vertebrae, number: 15 or fewer dorsal vertebrae. 167.1 Dorsal vertebrae, posterior dorsals, length of transverse processes: greater than centrum height. 195.0 Scapula, acromion process size: weakly developed. 273.1 Pubis, prepubic process, horizontal ridge on medial side: present. 275.0 Pubis, obturator foramen completely enclosed: present. |
| Ankylopollexia | 198.0 Scapula, deltoid ridge, orientation relative to long axis of scapula: close to parallel.
212.0 Humerus, deltopectoral crest form: a well-developed projection. 219.1 Ulna, flange on proximal end that wraps around the lateral edge of the radius: present. 227.1 Carpals, fusion: present. 232.1 Manus digit I, orientation: diverges at least 45 degrees from the antebrachial axis. 233.1 Metacarpal I, shape: short, block-like. 241.1 Manus ungual I, shape: subconical. 259.0 Ilium, brevis fossa, well defined by a lateral lip: absent. 323.1 Tendons, ossified epaxial and hypaxial tendon arrangement: double-layered lattice. |
| Styracosterna | 5.1 Premaxilla, oral margin denticulation: present.
123.2 Maxillary teeth, number: 18-28 positions. 137.1 Dentary teeth, maximum number of ridges extending from the base to the tip of the crown on lingual side of teeth: two to four ridges. 149.1 Maxillary and dentary crowns, relative width: maxillary crowns narrower. 164.1 Dorsal vertbrae, proportions of mid to posterior centra: length is much shorter than height. |
| Unnamed node | 180.1 Caudal vertebrae, shape of neural spines on proximal caudals: bowed caudally.
217.0 Ulna, length relative to dorsoventral thickness at mid-shaft: less than 9. 218.2 Ulna, olecranon process length as a percentage of total ulnar length: 17% or greater. 221.1 Radius, proportions: relatively robust, the minimal radial width is equal to or greater than 12% radial length. 223.1 Radius, tubercle near proximal end of radius for insertion of M. biceps: present. 242.1 Manus ungual I, length: greater than 30% the length of the radius. 276.1 Pubis, obturator foramen shape: circular. |
| Iguanodontidae | 42.1 Palpebral(s), length relative to rostrocaudal width of orbit if unfused to orbital rim: traverses entire width of orbit.
45.1 Postorbital, shape of squamosal process: bifurcated. 112.1 Surangular, position of small rostral fenestra: lies within the surangular. (Note that this fenestra is not present in Iguanodon or Equijubus.) |
| Mantellisaurus holotype + RBINS R57 | 15.1 Premaxilla-maxilla boundary, fossa or foramen just dorsal to the oral margin: present.
17.0 Maxilla, diastema length: one to two tooth widths. 95.1 Dentary ramus, shape of rostral end: strongly downturned. 105.1 Dentary, coronoid process orientation: subvertical. 164.0 Dorsal vertbrae, proportions of mid to posterior centra: craniocaudal length is subequal to or longer than dorsoventral height. 211.0 Humerus, length relative to femur: less than 62% of femoral length. 230.1 Metacarpal III, ratio between length and width at mid-shaft: long and slender, ratio greater than 5.5. |
| Bolong + Jinzhousaurus | 29.1 Antorbital fenestra: absent.
195.0 Scapula, acromion process size: weakly developed. 216.0 Humerus, relative width of medial and lateral distal condyles: lateral condyle wider. 217.0 Ulna, length relative to dorsoventral thickness at mid-shaft: less than 9. 221.1 Radius, proportions: relatively robust, the minimal radial width is equal to or greater than 12% radial length. 242.1 Manus ungual I, length: greater than 30% the length of the radius. 281.0 Ischium, shaft, shape in lateral view: straight. |
| Hadrosauroidea | 29.1 Antorbital fenestra: absent.
73.1 Squamosal, relationship of right and left squamosals on skull roof: separated by only a narrow band of the parietal. 93.1 Dentary, length of diastema: long, the width of three teeth or more. 94.1 Dentary, rostral extent of Meckel's groove meets the dentary symphysis: absent, ends more caudally. 103.2 Dentary: caudal extent of tooth row: terminates directly ventral to apex of the coronoid process or more caudally. 105.1 Dentary, coronoid process orientation: subvertical. 107.1 Coronoid process, rostrocaudally expanded apex: present. 136.0 Maxillary teeth, maximum number of ridges extending from the base to the tip of the crown on labial side of teeth: primary ridge only. 150.1 Dentary teeth, maximum number of functional teeth exposed on the occlusal plane: one functional tooth rostrally and caudally, and up to two teeth at and approaching the middle of the dental battery. 151.1 Dentary teeth, maximum number of replacement teeth: two. 152.1 Dentary alveoli: parallel grooves lining a continuous dental battery. 186.2 Chevrons, position of most proximal chevron: distal end of third caudal vertebra or more distal. 207.1 Coracoid, ratio between the length of the scapular articulation and the length of the lateral margin of the glenoid in lateral view: less than 1.25. 220.1 Radius, length: greater than 70% of humeral length. 230.1 Metacarpal III, ratio between length and width at mid-shaft: long and slender, ratio greater than 5.5. 251.0 Ilium, preacetabular process, morphology of distal end: parallel-sided or slightly tapering. 252.0 Ilium, cranial sacral facets transversely widened so that the base of the preacetabular process is thicker ventrally than dorsally: absent. 281.0 Ischium, shaft, shape in lateral view: straight. 305.1 Femur, distal condyles, shape of articular surface: rounded articular surface. |
