APPENDIX 3

List of morphological characters used in cladistic analysis. All characters taken directly from or modified from Wilson (2002). Character codings: (0), primitive state; (1, 2,...), derived states(s); (9), inapplicable character; (?), missing data.

1. Posterolateral processes of premaxilla and lateral processes of maxilla, shape: without midline contact (0); with midline contact, forming marked narial depression, subnarial foramen not visible laterally (1).

2. Premaxillary anterior margin, shape: without step (0); with marked step, anterior portion of skull sharply demarcated (1).

3. Maxillary border of external nares, length: short, making up much less than one-fourth narial perimeter (0); long, making up more than one-third narial perimeter (1).

4. Preantorbital fenestra: absent (0); present (1).

5. Subnarial foramen and anterior maxillary foramen, position: well distanced from one another (0); separated by narrow bony isthmus (1).

6. Antorbital fenestra, maximum diameter: much shorter than orbital maximum diameter (0); subequal to orbital maximum diameter (1).

7. Antorbital fossa: present (0); absent (1).

8. External nares, position: retracted to level of orbit (0); retracted to a position between orbits (1).

9. External nares, maximum diameter: shorter than orbital maximum diameter (0); longer than orbital maximum diameter (1).

10. Orbital ventral margin, anteroposterior length: broad, with subcircular orbital margin (0); reduced, with acute orbital margin (1).

11. Lacrimal, anterior process: present (0); absent (1).

12. Jugal-ectopterygoid contact: present (0); absent (1).

13. Jugal, contribution to antorbital fenestra: very reduced or absent (0); large, bordering approximately one-third its perimeter (1).

14. Prefontal, posterior process size: small, not projecting far posterior of frontal-nasal suture (0); elongate, approaching parietal (1).

15. Prefontal, posterior process shape: flat (0); hooked (1).

16. Postorbital, ventral process shape: transversely narrow (0); broader transversely than anteroposteriorly (1).

17. Postorbital, posterior process: present (0); absent (1).

18. Frontal contribution to supratemporal fossa: present (0); absent (1).

19. Frontals, midline contact (symphysis): sutured (0), or fused (1) in adult individuals.

20. Frontal, anteroposterior length: approximately twice (0) or less than (1) minimum transverse breadth.

21. Parietal occipital process, dorsoventral height: short, less than the diameter of the foramen magnum (0); deep, nearly twice the diameter of the foramen magnum (1).

22. Parietal, contribution to post-temporal fenestra: present (0); absent (1).

23. Postparietal foramen: absent (0); present (1).

24. Parietal, distance separating supratemporal fenestrae: less than the long axis of the supratemporal fenestra (0); twice the long axis of the supratemporal fenestra (1).

25. Supratemporal fenestra: present (0); absent (1).

26. Supratemporal fenestra, long axis orientation: anteroposterior (0); transverse (1).

27. Supratemporal fenestra, maximum diameter: much longer than (0) or subequal to (1) that of foramen magnum.

28. Supratemporal region, anteroposterior length: temporal bar longer (0) or shorter (1) anteroposteriorly than transversely.

29. Supratemporal fossa, lateral exposure: not visible laterally, obscured by temporal bar (0); visible laterally, temporal bar shifted ventrally (1).

30. Laterotemporal fenestra, anterior extension: posterior to orbit (0); ventral to orbit (1).

31. Squamosal-quadratojugal contact: present (0); absent (1).

32. Quadratojugal, anterior process length: short, anterior process shorter than dorsal process (0); long, anterior process more than twice as long as dorsal process (1).

33. Quadrate fossa: absent (0); present (1).

34. Quadrate fossa, depth: shallow (0); deeply invaginated (1).

35. Quadrate fossa, orientation: posterior (0); posterolateral (1).

36. Palatobasal contact, shape: pterygoid with small facet for basipterygoid articulation (0); dorsomedailly oriented hook for basipterygoid articulation (1); rocker-like surface for basipterygoid articulation (2).

37. Pterygoid, transverse flange (i.e., ectopterygoid process) position: between orbit and antorbital fenestra (0); anterior to antorbital fenestra (1).

38. Pterygoid, quadrate flange size: large, palatobasal and quadrate articulations well separated (0); small, palatobasal and quadrate articulations approach (1).

39. Pterygoid, palatine ramus shape: straight, at level of dorsal margin of quadrate ramus (0); stepped, raised above level of quadrate ramus (1).

40. Palatine, lateral ramus shape: plate-shaped (long maxillary contact) (0); rod-shaped (narrow maxillary contact) (1).

41. Epipterygoid: present (0); absent (1).

42. Vomer, anterior articulation: maxilla (0); premaxilla (1).

43. Supraoccipital, height: twice (0) subequal to or less than (1) height of foramen magnum.

44. Paroccipital process, ventral nonarticular process: absent (0); present (1).

45. Crista prootica, size: rudimentary (0); expanded laterally into `dorsoventral process' (1).

46. Basipterygoid processes, length: short, approximately twice basal diameter (0); elongate, at least four times basal diameter (1).

47. Basipterygoid processes, angle of divergence: approximately 45° (0); less than 30° (1).

48. Basal tubera, anteroposterior depth: approximately half dorsovenral height (0); sheet-like, 20% dorsoventral height (1).

49. Basal tubera, breadth: much broader than (0) or narrower than (1) occipital condyle.

50. Basioccipital depression between foramen magnum and basal tubera: absent (0); present (1).

51. Basisphenoid/basipterygoid recess: present (0); absent (1).

52. Basisphenoid-quadrate contact: absent (0); present (1).

53. Basipterygoid processes, orientation: perpendicular to skull roof (0); angled approximately 45° to skull roof (1).

54. Occipital region of skull, shape: anteroposteriorly deep, paroccipital processes oriented posterolaterally (0); flat, paroccipital processes oriented transversely (1).

55. Dentary, depth of anterior end of ramus: slightly less than that of dentary at midlength (0); 150% minimum depth (1).

56. Dentary, anteroventral margin shape: gently rounded (0); sharply projecting triangular process or 'chin' (1).

57. Dentary symphysis, orientation: angled 15° or more anteriorly to (0) or perpendicular to (1) axis of jaw ramus.

58. External mandibular fenestra: present (0); absent (1).

59. Surangular depth: less than twice maximum depth of the angular (0) more than two and one-half times maximum depth of the angular (1).

60. Surangular ridge separating adductor and articular fossae: absent (0); present (1).

61. Adductor fossa, medial wall depth: shallow (0); deep, prearticular expanded dorsoventrally (1).

62. Splenial posterior process, position: overlapping angular (0); separating anterior portions of prearticular and angular (1).

63. Splenial posterodorsal process: present, approaching margin of adductor chamber (0); absent (1).

64. Coronoid, size: extending to dorsal margin of jaw (0); reduced, not extending dorsal to splenial (1); absent (2).

65. Tooth rows, shape of anterior portions: narrowly arched, anterior portion of tooth rows V-shaped (0); broadly arched, anterior portion of tooth rows U-shaped (1); rectangular, tooth-bearing portion of jaw perpendicular to jaw rami (2).

66. Tooth rows, length: restricted anterior to orbit (0); restricted anterior to subnarial foramen (1).

67. Crown-to-crown occlusion: absent (0); present (1).

68. Occlusal pattern: interlocking, V-shaped wear facets (0); high-angled planar facets (1); low-angled planar facets (2).

69. Tooth crowns, orientation: aligned along jaw axis, crowns do not overlap (0); aligned slightly anterolingually, tooth crowns overlap (1).

70. Tooth crowns, cross-sectional shape at mid-crown: elliptical (0); D-shaped (1); cylindrical (2).

71. Enamel surface texture: smooth (0); wrinkled (1).

72. Marginal tooth denticles: present (0); absent on posterior edge (1); absent on both anterior and posterior edges (2).

73. Dentary teeth, number: greater than 20 (0); 17 or fewer (1).

74. Replacement teeth per alveolus, number: two or fewer (0); more than four (1).

75. Teeth, orientation: perpendicular to jaw margin (0); oriented anteriorly relative to jaw margin (1).

76. Teeth, longitudinal grooves on lingual aspect: absent (0); present (1).

77. Presacral vertebrae, bone texture: solid (0); spongy, with large, open internal cells, 'camellate' (1).

78. Presacral centra, pleurocoels: absent (0); present (1).

79. Atlantal intercentrum, occipital facet shape: rectangular in lateral view, length of dorsal aspect subequal to that of ventral aspect (0); expanded anteroventrally in lateral view, anteroposterior length of dorsal aspect shorter than that of ventral aspect (1).

80. Cervical vertebrae, number: 9 or fewer (0); 10 (1); 12 (2); 13 (3); 15 or greater (4).

81. Cervical neural arch lamination: well developed, with well defined laminae and coels (0); rudimentary, diapophyseal laminae only feebly developed if present (1).

82. Cervial centra, articular face morphology: amphicoelous (0); opisthocoelous (1).

83. Cervical pleurocoels, shape: simple, undivided (0); complex, divided by bony septa (1).

84. Anterior cervical centra, height:width ratio: less than 1 (0); approximately 1.25 (1).

85. Anterior cervical neural spines, shape: single (0); bifid (1).

86. Mid-cervical centra, anteroposterior length/height of posterior face: 2.5-3.5 (0); greater than 4.0 (1).

87. Mid-cervical neural arches, height: less than that of posterior centrum face (0); greater than that of posterior centrum face (1).

88. Middle and posterior cervical neural arches, centroprezygapophyseal lamina (cprl), shape: single (0); divided (1).

89. Posterior cervical and anterior dorsal neural spines, shape: single (0); bifid (1).

90. Posterior cervical and anterior dorsal bifid neural spines, median tubercle: absent (0); present (1).

91. Dorsal vertebrae, number: 15 (0); 14 (1); 13 (2); 12 (3); 11 (4); 10 or fewer (5).

92. Dorsal neural spines, breadth: narrower (0) or much broader (1) transversely than anteroposteriorly.

93. Dorsal neural spines, length: approximately twice (0) or approximately four times (1) centrum length.

94. Anterior dorsal centra, articular face shape: amphicoelous (0); opisthocoelous (1).

95. Middle and posterior dorsal neural arches, centropostzygapophyseal lamina (cpol), shape: single (0); divided (1).

96. Middle and posterior dorsal neural arches, anterior centroparapophyseal lamina (acpl): absent (0); present (1).

97. Middle and posterior dorsal neural arches, prezygoparapophyseal lamina (prpl): absent (0); present (1).

98. Middle and posterior dorsal neural arches, posterior centroparapophyseal lamina (pcpl): absent (0); present (1).

99. Middle and posterior dorsal neural arches, spinodiapophyseal lamina (spdl): absent (0); present (1).

100. Middle and posterior dorsal neural arches, spinopostzygapophyseal lamina (spol), shape: single (0); divided (1).

101. Middle and posterior dorsal neural arches, spdl and spol contact: absent (0); present (1).

102. Middle and posterior dorsal neural spines, shape: tapering or not flaring distally (0); flared distally, with pendant, triangular lateral processes (1).

103. Middle and posterior dorsal neural arches, `infradiapophyseal' pneumatopore between scdl and pcdl: absent (0); present (1).

104. Middle and posterior dorsal neural spines, orientation: vertical (0); posterior, neural spine summit approaches level of diapophyses (1).

105. Posterior dorsal centra, articular face shape: amphicoelous (0); opisthocoelous (1).

106. Posterior dorsal neural arches, hyposphene-hypantrum articulations: present (0); absent (1).

107. Posterior dorsal neural spines, shape: rectangular through most of length (0); `petal' shaped, expanding transversely through 75% of its length and then tapering (1).

108. Sacral vertebrae, number: 3 or fewer (0); 4 (1); 5 (2); 6 (3).

109. Sacrum, sacricostal yoke: absent (0); present (1).

110. Sacral neural spines, length: approximately twice length of centrum (0); four times length of centrum (1).

111. Sacral ribs, dorsoventral length: low, not projecting beyond dorsal margin of ilium (0); high, extending beyond dorsal margin of ilium (1).

112. Caudal bone texture: solid (0); spongy, with large internal cells (1).

113. Caudal vertebrae, number: more than 45 (0); 35 or fewer (1).

114. Caudal transverse processes: persist through caudal 20 or more posteriorly (0); disappear by caudal 15 (1); disappear by caudal 10 (2).

115. First caudal centrum, articular face shape: flat (0), procoelous (1), opisthocoelous (2); biconvex (3).

116. First caudal neural arch, coel on lateral aspect of neural spine: absent (0); present (1).

117. Anterior caudal centra (excluding the first), articular face shape: amphiplatyan or platycoelous (0); strongly procoelous (1); opisthocoelous (2).

118. Anterior caudal centra, pleurocoels: absent (0), present (1).

119. Anterior caudal centra, length: approximately the same over the first 20 vertebrae (0); doubling over the first 20 vertebrae (1).

120. Anterior caudal neural arches, spinoprezygapophyseal lamina (sprl): absent (0); present and extending onto lateral aspect of neural spine (1).

121. Anterior caudal neural arches, sprl-spol contact: absent (0); present, forming a prominent lamina on lateral aspect of neural spine (1).

122. Anterior caudal neural arches, prespinal lamina (prsl): absent (0); present (1).

123. Anterior caudal neural arches, postspinal lamina (posl): absent (0); present (1).

124. Anterior caudal neural arches, postspinal fossa: absent (0); present (1).

125. Anterior caudal neural spines, transverse breadth: approximately 50% of anteroposterior length (0); greater than anteroposterior length (1).

126. Anterior caudal transverse processes, proximal depth: shallow, on centrum only (0); deep, extending from centrum to neural arch (1).

127. Anterior caudal transverse processes, shape: triangular, tapering distally (0); 'wing-like', not tapering distally (1).

128. Anterior caudal transverse processes, diapophyseal laminae (acdl, pcdl, prdl, podl): some absent (0); all present (1).

129. Anterior caudal transverse processes, acdl shape: single (0); divided (1).

130. Anterior and middle caudal centra, shape: cylindrical (0); quadrangular, flat ventrally and laterally (1).

131. Anterior and middle caudal centra, ventral longitudinal hollow: absent (0); present (1).

132. Middle caudal neural spines, orientation: angled posterodorsally (0), vertical (1), angled anterodorsally (2).

133. Middle and posterior caudal centra, anterior articular face shape: flat (0); procoelous (cone shaped) (1); opisthocoelous (2).

134. Posterior caudal centra, shape: cylindrical (0); dorsoventrally flattened, breadth at least twice height (1).

135. Distalmost caudal centra, articular face shape: platycoelous (0); biconvex (1).

136. Distalmost biconvex caudal centra, length-to-height ratio: less than 4 (0); greater than 5 (1).

137. Distalmost biconvex caudal centra, number: 10 or fewer (0); more than 30 (1).

138. Cervical rib, tuberculum-capitulum angle: greater than 90° (0); less than 90°, rib ventrolateral to centrum (1).

139. Cervical ribs, length: much longer than centrum, overlapping as many as three subsequent vertebrae (0); shorter than centrum, little or no overlap (1).

140. Dorsal ribs, proximal pneumatocoels: absent (0); present (1).

141. Anterior dorsal ribs, cross-sectional shape: sub-circular (0); plank-like, anteroposterior breadth more than three times mediolateral breadth (1).

142. 'Forked' chevrons with anterior and posterior projections: absent (0); present (1).

143. 'Forked' chevrons, distribution: distal tail only (0); throughout middle and posterior caudal vertebrae (1).

144. Chevrons, 'crus' bridging dorsal margin of haemel canal: present (0); absent (1).

145. Chevron haemal canal, depth: short, approximately 25% chevron length (0); long, approximately 50% chevron length (1).

146. Chevrons: persisting throughout at least 80% of tail (0); disappearing by caudal 30 (1).

147. Posterior chevrons, distal contact: fused (0); unfused (open) (1).

148. Posture: bipedal (0); columnar, obligately quadrupedal posture (1).

149. Scapular acromion process, size: narrow (0); broad, width more than 150% minimum width of blade (1).

150. Scapular blade, orientation: forming close to a 90° angle with coracoid articulation (0); forming a 45° angle with coracoid articulation (1).

151. Distal scapular blade, shape: acromial edge not expanded (0); acromial edge expanded (1); racquet-shaped (2).

152. Scapular glenoid, orientation: relatively flat or laterally facing (0); strongly bevelled medially (1).

153. Scapular blade, cross-sectional shape: flat or rectangular (0); D-shaped (1).

154. Coracoid, proximodistal length: less than length of scapular articulation (0) approximately twice length of scapular articulation (1).

155. Coracoid, anteroventral margin shape: rounded (0); rectangular (1).

156. Coracoid, infraglenoid lip: absent (0); present (1).

157. Sternal plate, shape: oval (0); crescentic (1).

158. Humeral proximolateral corner, shape: rounded (0); square (1).

159. Humeral deltopectoral attachment, development: prominent (0); reduced to a low crest or ridge (1).

160. Humeral deltopectoral crest, shape: relatively narrow throughout length (0); markedly expanded distally (1).

161. Humeral midshaft cross-section, shape: circular (0); elliptical, with long axis oriented transversely (1).

162. Humeral distal condyles, articular surface shape: restricted to distal portion of humerus (0); exposed on anterior portion of humeral shaft (1).

163. Humeral distal condyle, shape: divided (0); flat (1).

164. Ulnar proximal condyle, shape: subtriangular (0); triradiate, with deep radial fossa (1).

165. Ulnar proximal condylar processes, relative lengths: subequal (0); unequal, anteromedial arm longer (1).

166. Ulnar olecranon process, development: prominent, projecting above proximal articulation (0); rudimentary, level with proximal articulation (1).

167. Ulna, length-to-proximal breadth ratio: gracile (0); stout (1).

168. Radial distal condyle, shape: round (0); subrectangular, flattened posteriorly and articulating in front of ulna (1).

169. Radius, distal condyle orientation: perpendicular to long axis of shaft (0); bevelled approximately 20° proximolaterally relative to long axis of shaft (1).

170. Humerus-to-femur ratio: less than 0.60 (0); 0.60 or more (1).

171. Carpal bones, number: 3 or more (0); 2 or fewer (1).

172. Carpal bones, shape: round (0); block-shaped, with flattened proximal and distal surfaces (1).

173. Metacarpus, shape: spreading (0); bound, with subparallel shafts and articular surfaces that extend half their length (1).

174. Metacarpals, shape of proximal surface in articulation: gently curving, forming a 90° arc (0); U-shaped, subtending a 270° arc (1).

175. Longest metacarpal-to-radius ratio: close to 0.3 (0); 0.45 or more (1).

176. Metacarpal I, length: shorter than metacarpal IV (0); longer than metacarpal IV (1).

177. Metacarpal I, distal condyle shape: divided (0); undivided (1).

178. Metacarpal I distal condyle, transverse axis orientation: bevelled approximately 20° proximodistally with respect to axis of shaft (0); perpendicular to axis of shaft (1).

179. Manual digits II and III, phalangeal number: 2-3-4-3-2 or more (0); reduced, 2-2-2-2-2 or less (1); absent or unossified (1).

180. Manual phalanx I.1, shape: rectangular (0); wedge-shaped (1).

181. Manual nonungual phalanges, shape: longer proximodistally than broad transversely (0); broader transversely than long proximodistally (1).

182. Pelvis, anterior breadth: narrow, ilia longer anteroposteriorly than distance separating preacetabular processes (0); broad, distance bewteen preacetabular processes exceeds anteroposterior length of ilia (1).

183. Ilium, ischial peduncle size: large, prominent (0); low, rounded (1).

184. Iliac blade dorsal margin, shape: flat (0); semicircular (1).

185. Iliac preacetabular process, orientation: anterolateral to body axis (0); perpendicular to body axis (1).

186. Pubis, ambiens process development: small, confluent with anterior margin of pubis (0); prominent, projecting anteriorly from anterior margin of pubis (1).

187. Pubic apron, shape: flat (straight symphysis) (0); canted anteromedially (gentle S-shaped symphysis) (1).

188. Puboischial contact, length: approximately one-third total length of pubis (0); approximately one-half total length of pubis (1).

189. Ischial shaft, length: much shorter than shaft of pubis (0); equal to or longer than shaft of pubis (1).

190. Ischial shaft, shape: emarginate distal to pubic peduncle (0); no emargination distal to pubic peduncle (1).

191. Ischial distal shaft, shape: triangular, depth of ischial shaft increases medially (0); blade-like, medial and lateral depths subequal (1).

192. Ischial distal shafts, cross-sectional shape: V-shaped, forming an angle of nearly 50° with each other (0); flat, nearly coplanar (1).

193. Femoral fourth trochanter, development: prominent (0); reduced to crest or ridge (1).

194. Femoral lesser trochanter: present (0); absent (1).

195. Femoral mid-shaft, transverse diameter: subequal to anteroposterior diameter (0); 125-150% anteroposterior diameter (1); at least 185% anteroposterior diameter (2).

196. Femoral shaft, lateral margin shape: straight (0); proximal one-third deflected medially (1).

197. Femoral distal condyles, relative transverse breadth: subequal (0); tibial much broader than fibular (1).

198. Femoral distal condyles, orientation: perpendicular or slightly bevelled dorsolaterally relative to femoral shaft (0); bevelled dorsomedially approximately 10° relative to femoral shaft (1).

199. Femoral distal condyles, articular face shape: restricted to distal portion of femur (0); expanded onto anterior portion of femoral shaft (1).

200. Tibial proximal condyle, shape: narrow, long axis anteroposterior (0), expanded transveresly, condyle subcircular (1).

201. Tibial cnemial crest, orientation: projecting anteriorly (0); projecting laterally (1).

202. Tibia, distal breadth: approximately 125% mid-shaft breadth (0); more than twice mid-shaft breadth (1).

203. Tibial distal posteroventral process, size: broad transversely, covering posterior fossa of astragalus (0); shortened transversely, posterior fossa of astragalus visible posteriorly (1).

204. Fibula, proximal tibial scar, development: not well-marked (0); well-marked and deepening anteriorly (1).

205. Fibula, lateral trochanter: absent (0); present (1).

206. Astragalus, shape: rectangular (0); wedge-shaped, with reduced anteromedial corner (1).

207. Astragalus, foramina at base of ascending process: present (0); absent (1).

208. Astragalus, ascending process length: limited to anterior two-thirds of astragalus (0); extending to posterior margin of astragalus (1).

209. Astragalus, posterior fossa, shape: undivided (0); divided by vertical crest (1).

210. Astragalus, transverse length: 50% more than proximodistal height (0); subequal to proximodistal height (1).

211. Calcaneum: present (0); absent or unossified (1).

212. Distal tarsals 3 and 4: present (0); absent or unossified (1).

213. Metatarsus, posture: bound (0); spreading (1).

214. Metatarsal I proximal condyle, transverse axis orientation: perpendicular to axis of shaft (0); angled ventromedially approximately 15° to axis of shaft (1).

215. Metatarsal I distal condyle, transverse axis orientation: perpendicular to axis of shaft (0); angled dorsomedially to axis of shaft (1).

216. Metatarsal I distal condyle, posterolateral projection: absent (0); present (1).

217. Metatarsal I, minimum shaft width: less than that of metatarsals II-IV (0); greater than that of metatarsals II-IV (1).

218. Metatarsal I and V proximal condyle, size: smaller than those of metatarsals II and IV (0); subequal to those of metatarsals II and IV (1).

219. Metatarsal III, length: more than 30% that of tibia (0); less than 25% that of tibia (1).

220. Metatarsals III and IV, minimum transverse shaft diameters: subequal to that of metatarsals I or II (0); less than 65% that of metatarsals I or II (1).

221. Metatarsal V, length: shorter than length of metatarsal IV (0); at least 70% length of metatarsal IV (1).

222. Pedal nonungual phalanges, shape: longer proximodistally than broad transversely (0); broader transversely than long proximodistally (1).

223. Pedal digits II-IV, penultimate phalanges, development: subequal in size to more proximal phalanges (0); rudimentary or absent (1).

224. Pedal unguals, orientation: aligned with digit axis (0); deflected lateral to digit axis (1).

225. Pedal digit I ungual, length relative to pedal digit II ungual: subequal (0); 25% larger than that of digit II (1).

226. Pedal digit I ungual, length: shorter than metatarsal I (0); longer than metatarsal I (1).

227. Pedal ungual I, shape: broader transversely than dorsoventrally (0); sickle-shaped, much deeper dorsoventrally than broad transversely (1).

228. Pedal ungual II-III, shape: broader transversely than dorsoventrally (0); sickle-shaped, much deeper dorsoventrally than broad transversely (1).

229. Pedal digit IV ungual, development: subequal in size to unguals of pedal digits II and III (0); rudimentary or absent (1).

230. Osteoderms: absent (0); present (1).

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