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Appendix 2.1. Taxon-Character Matrix

Appendix 2.2. Character Choice and State Definitions for the Phylogenetic Analysis Conducted in This Study

1. Premaxillary rostrum: dorsal aspect. Margins converge to a blunt tip (0), modest rounded expansion (1), occlusal margin is broad and rounded in dorsal view such that its overall width approaches that of the skull roof (2), flared occlusal margins that form a “spoon-bill” structure in dorsal view (3).

2. Premaxilla: level of occlusal margin relative to that of the maxillary tooth row. Not at all (or slightly) ventrally offset from alveolar margin of the maxilla (0), strongly ventrally offset (1).

3. Premaxillary teeth: present (0), absent (1).

4. Premaxillary denticles: absent (0), present (1).

5. External naris: confined to area above oral margin of premaxilla (0), posterior margin extended posteriorly to lie above the maxilla (1).

6. Premaxilla: anterolateral margin of the nasal fossa, above the occlusal edge of the premaxilla is reflected dorsally to form a distinct rim: absent (0), present (1).

7. Premaxilla-lacrimal contact: absent (0), present (1) posterolateral premaxillary process extends posterodorsally to also contact/overlap the prefrontal (2).

8. Premaxillary posterolateral process: tapers to point (0), posterior tip is bluntly truncated (1).

9. Antorbital fenestra shape. Opening when viewed laterally: large and subtriangular (0), small and subcircular (1), absent (2).

10. Antorbital fenestra location: between lacrimal and maxilla (0), on anterodorsal margin (premaxillary suture) of maxilla and not visible in lateral view of the articulated skull (1).

11. Lacrimal-nasal contact: present (0), absent (1).

12. Maxilla: dorsal process morphology. Flattened mound-like structure (0), narrow, finger-like process (1), laterally flattened subtriangular plate (2).

13. Jugal: anterior process. Tapering (0), expanded (1), expanded and truncated anteriorly (2).

14. Jugal-maxilla suture: scarf joint (0), “finger-in-recess” (oblique finger-like process of the maxilla fits into a slot formed in the medioventral surface of the anterior jugal ramus) (1), butt-jointed against a broad facet on the lateral surface of the ascending process of the maxilla (2).

15. Jugal and its free ventral margin: generally strap-like with little undulation along the ventral edge (0), marked sinuous ventral edge with marked ventral deflection posteriorly (1).

16. Jugal-ectopterygoid contact: present (0), absent (1). Difficult to score in many instances.

17. Jugal contribution to the margin of the infratemporal fenestra: jugal forms part of the margin (0), jugal forms the entire margin by excluding the quadratojugal (1).

18. Quadratojugal foramen: absent (0), present (1). Very limited distribution.

19. Quadrate (paraquadratic) foramen: between quadratojugal and quadrate. Present (0), absent (1).

20. Quadrate: embayment on anterolateral wing. Relatively small and semicircular in outline (0), broad embayment whose edges are marked by a scarf suture for a close-fitting quadratojugal (1).

21. Quadrate: posterior margin of the shaft. Bowed anteriorly and the dorsal portion tilted posteriorly (0), straight (1). Often difficult to score because of postmortem distortion.

22. Quadrate-articular condyle: transversely expanded (0), laterally compressed so that it forms (almost) a simple rounded condyle (1).

23. Palpebral (supraorbital) bone(s): present (0), absent or fused to orbital margin (1). Given the looseness of attachment of this bone to the orbital margin (and therefore the lack of a reliable osteological marker) evidence of absence is subjective.

24. Frontal: shape. Arched and narrow embayed laterally so that the orbital cavity is exposed dorsally (0), flat and transversely broad plate, roofing the orbital cavity dorsally (1), anteroposteriorly abbreviated (2).

25. Frontal: forms part of the dorsal margin of the orbital cavity. Present (0), absent (1).

26. Postorbital-squamosal contact: postorbital forms a tapering finger-like squamosal process the overlaps the squamosal (0), this process develops a bifurcate tip (1). McDonald et al. (2010) proposed that this process might be coded in three ways: “blunt,” “pointed,” or “bifurcate.” However, there is variation in this structure as it is illustrated in skull reconstructions, which makes the distinction between “blunt” and “pointed” one that is potentially subjective and risks misinterpretation.

27. Foramen magnum dorsal margin: supraoccipital in dorsal margin (0), supraoccipital excluded from dorsal margin by exoccipitals (1).

28. Foramen magnum ventral margin: basioccipital in ventral margin (0), basioccipital excluded from ventral margin by the exoccipitals (1). This character, though commonly used and perhaps of more value at a coarser scale (e.g., across higher-level taxonomic groupings), is often difficult to assess reliably (within individual taxa), and may vary ontogenetically and be subject by taphonomic influence.

29. Paroccipital wing shape: horizontal and dorsoventrally expanded distally (0), pendant distal tip (1).

30. Basipterygoid process orientation: anteroventral (0), posterolateral (1).

31. Predentary occlusal margin: smooth edged (0), denticulate (1).

32. Predentary shape (in plan view): subtriangular (0), arcuate (1), broad and subrectangular (2).

33. Predentary ventral lobe: median tab sometimes notched in the midline (0), deeply incised in the midline and with strongly bifurcate lobes (1).

34. Predentary rostral surface: smooth (0); bearing a pair of oblique grooves, one on either side of midline (1), midline groove (2).

35. Mandibular diastema (the gap between the posterior end of the predentary and the first dentary alveolus): absent (0), present (1), greater than three crown widths (2). Difficult to assess in instances in which the predentary is not articulated with the dentary and/or when the anterior part of the dentary is not well preserved.

36. Dentary ramus shape in lateral view: straight (0), arched along its ventral edge (1).

37. Dentary ramus (tooth-bearing portion) shape: tapers anteriorly (0), parallel dorsal and ventral borders (1), deepens anteriorly (2).

38. Dentary coronoid process profile: axis of coronoid oblique (posterodorsal orientation) (0), perpendicular (1), anteriorly inclined (2).

39. Dentary coronoid process shape: dorsal tip unexpanded (0), expanded (1).

40. Dentary coronoid process position: laterally offset and dentition [alveoli] curves laterally into its base (0), posterior dentition extends medial to the middle of the coronoid process (1), posterior dentition extends posterior to the coronoid process (2).

41. Surangular foramen: present (0), absent (1).

42. Surangular-angular suture: obliquely inclined (0), horizontal (1). Difficult to score faithfully because this feature may be based upon skull reconstructions which may, or may not, be accurate in this area.

43. Angular: lateral exposure. Visible laterally (0), not visible (contact with surangular is positioned ventrally or medially) (1).

44. Replacement crowns present: one (0), two (1), three or more (2).

45. Wear facet distribution on dentary and maxillary crowns: irregular and discontinuous distribution on individual crowns (0), wear facets continuous across adjacent crowns, producing a uniformly narrow cutting/grinding surface (1), the oldest and successional crowns contribute to the wear surface to varying degrees to produce a transversely broader cutting/grinding occlusal (2).

46. Relative crown width: maxillary crowns equal in width to dentary crowns (0), narrower than dentary crowns (1), equal in width to dentary crowns, but “miniaturized” (2).

47. Enamel surface distribution on tooth crowns: equally distributed on both sides of crown (0), asymmetrical distribution, thicker on one surface of the crown (1), enamel is restricted to one side of the crown (2).

48. Marginal denticle shape: simple cones (0), tongue-shaped (1), curved ledges with mammillae (2), absent, or reduced to small irregular papillae (3).

49. Tooth roots: tapering cylinders (0), longitudinally grooved to accommodate adjacent, closely packed teeth (1), highly angular sided “prismatic” roots (2).

50. Dentary tooth curvature of long axis: the root–crown axis of the tooth is straight (0), the long axis of the tooth is bowed lingually so that the occlusal portion of the crown becomes directed labiodorsally (1).

51. Dentary teeth, crown shape in lingual view: simple, symmetrical leaf-shaped profile (0), broad, shield-like (1), in unworn examples occlusal margin forms a distinct shoulder mesially (2), mesiodistally compressed and diamond shaped (3).

52. Dentary teeth, presence of oblique, thickened ledges at the base of the enamelled lingual face of the crown: absent (0), present (1).

53. Dentary teeth, primary ridge: absent (0), median position (1), distally offset (2).

54. Dentary teeth, ridge pattern: simple median swelling (0), prominent primary ridge with variable number of parallel subsidiary ridges (1), parallel primary and secondary ridge divide crown face into three zones (2), primary ridge alone (3).

55. Dentary teeth, relative size in mandible: large and shield shaped (0), miniaturized (1).

56. Dentary, lateral alveolar wall shape: shaped by dentary crowns (0), narrow, parallel-sided grooves (1).

57. Maxillary teeth shape 1: equal width to dentary crowns (0), narrower and more lanceolate than opposing dentary crowns (1), lanceolate but equal in width to dentary crowns (2).

58. Maxillary teeth shape 2: Root–crown long axis of the tooth straight (0); long axis bowed, convex labially (1).

59. Maxillary teeth, labial ridges 1: simple median swelling (0), very prominent primary ridge (1), several subsidiary ridges and no obvious primary ridge (2).

60. Maxillary teeth, labial ridges 2: simple medial swelling (0), array of primary, secondary, and subsidiary ridges (1), single median primary ridge with no other ridges present (2).

61. Axis vertebra. Neural spine shape: low and sloping (0), tall and expanded (1).

62. Cervical vertebrae. Centrum articular surfaces: amphiplatyan (0), opisthocoelous (1).

63. Dorsal vertebrae. Neural spine shape: low and rectangular (0), tall and narrow (1). Extremely tall, relative to the anteroposterior dimension of spine (2). Dubious. Biomechanical: small animals have short neural spines. Only two taxa, Ouranosaurus and Hypselospinus, have extremely elongate neural spines—i.e., probable homoplasy.

64. Epaxial ossified tendons: arranged in linear bundles (0), form a layered lattice against the neural spines (1). Dubious: reflects the accommodation space available: taller spines enable a lattice-like arrange of tendons to exist.

65. Ossified tendons form a posterior caudal sheath: absent (0), present (1). Rarely preserved, and where a sheath of tendons is known, its occurrence seems to be restricted to basal clypeodontans and may be plesiomorphic within cerapodans.

66. Scapular acromion: prominent on the proximodorsal margin of scapula (0), occupying a median position on the proximal shaft and curved toward the dorsal edge of the proximal scapular blade (1), developed into a promontory that overhangs the proximal end of the scapula and is not curved toward the dorsal border (2).

67. Sternal shape: reniform (0), hatchet shaped with a stout, short “handle” (1), pronounced elongation of the “handle” of the hatchet relative to the “blade” (2).

68. Carpals: fully ossified (0), fused to form a carpometacarpal I block (1), reduced to no more than two small ossicles (2).

69. Metacarpal I: elongate dumbbell shaped (0), short, block-like and fused against carpals (1), absent (2).

70. Metacarpals II–IV: spreading (0), robust, closely appressed (1), slender and elongate (2).

71. Manus digit I: present (0), absent (1).

72. Ungual of manus digit I: claw-like (0), subconical (1), lost (2).

73. Unguals of manus digits II and III: claw-like (0), flattened, twisted and hoof-like (1).

74. Manus digit III: four phalanges (0), three phalanges (1).

75. Ilium, preacetabular process: long, laterally compressed (0), axially twisted so that lateral surface faces dorsolaterally (1).

76. Ilium: profile of dorsal edge. Horizontal to slightly arched, no significant notch in its profile posterodorsal to the ischiadic peduncle (0), sinuous profile created by the presence of a broad saddle-like notch (1).

77. Ilium: dorsal margin development: no transverse thickening of the dorsal edge in the region posterodorsal to the ischial peduncle (0), transversely thickened, beveled edge (1), thickened edge developed into a thick rolled edge (2), prominently everted, with a downturned, flap-shaped pendule that overhangs the ischiadic peduncle (3).

78. Ilium: postacetabular process in profile. Vertical plate with rounded edge (0), generally triangular, tapering posteriorly (1), laterally compressed and relatively narrow, rectangular bar (2), upturned plate (3).

79. Pubis: anterior ramus form: short (0), elongate (1).

80. Pubis: anterior ramus shape. Short, deep and blunt (0), rod-shaped (1), laterally compressed parallel-sided blade (2), expanded distally (3), strongly transversely compressed and deeply expanded distal portion (4).

81. Pubis: posterior ramus. Terminates bluntly adjacent to distal end of ischium (0), shorter than ischium, tapers to a point (1).

82. Ischium: shaft morphology 1. Straight (0), bowed (1).

83. Ischium: shaft morphology 2. Compressed and blade-like along length of shaft (0), sub-cylindrical shaft (1), narrow, angular-sided shaft (2).

84. Ischium: shaft morphology 3. Distal end unexpanded (0), distal end expanded into “boot” (1), distal end laterally expanded, rather than expanded anteroposteriorly (2).

85. Ischium: obturator process. Absent (0), positioned near mid-shaft (1), positioned close to pubic peduncle, from which it is separated by an embayment (2).

86. Femoral head grooved posteriorly: present (0), absent (1).

87. Femur: curvature of shaft. Femoral shaft bowed along its length (0), proximal half of the shaft straight, distal half of shaft curved caudally (1), femoral shaft straight (2).

88. Femur: fourth trochanter. Pendant (0), large, triangular (1), curved, laterally compressed eminence (2).

89. Femur: distal extensor groove. Absent (0), very broad trough (1), U-shaped trough (2), partially enclosed by expansion of adjacent anterior condyles (3), edges of trough meet to form a fully-enclosed tunnel (4).

90. Femur: distal condyles. Moderately expanded anteroposteriorly (0), strongly expanded and partly occluding flexor channel (1). Most probably a size-related mechanical feature.

91. Metatarsal I: well developed and articulates with proximal phalanx (0), slender and splint-like (1), absent (2). Difficult to assess in many instances. As in the case of a palpebral bone on the orbital margin, is absence actual evidence of absence, or a biostratinomic artifact?

92. Pedal phalanges: shape. Dorsoventrally flattened but elongate and pointed (0), elongate, bluntly truncated tip with prominent claw grooves retained (1), anterior margin broadly rounded in dorsal view, lateral claw grooves absent (2).