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SUPPLEMENTAL DATA 1

Here is a list of things which were changed by OM on the digital Irritator skull reconstruction to make it mostly symmetrical and articulate properly. 

Where bones seemed to be mostly intact, OM opted to not deform those bones and instead focused on making other, less well preserved or obviously deformed bones, fit. OM did not include repositioning of bones in this list.

- Restoring the fossils original shape before breaking

The original fossil was broken in two at one point and has been incorrectly glued back together. The whole anterior part of the skull was rotated slightly to the left side. OM put this back in place, rotating it to the right, using all the bones that were present in both the anterior and posterior half of the fossil. Aligning this, fixed bending in the nasals, maxillae, palatines, pterygoids and surangulars.

- Bending of the nasals and adjacent bones

Lateral bending of the nasals in two main places. As mentioned above, the fossil had a bend at the break of the fossil, so OM corrected for the lateral shift and rotated the anterior snout piece slightly to the right. The second location is just behind the posterordorsal processes of the maxillae. This is probably where the nasals and lacrimals are deformed the most. OM bent the nasals and anterodorsal processes of the lacrimals to the left. These two changes made the nasals roughly straight.

- Bending and shifting of the maxillae

Much of the skull was pushed in medially on the right side. This effectively collapsed much of the skull laterally. As a result, the posterior half of both maxillae was laterally pushed to the left. OM corrected this, making them more symmetrical. Around the mid-point of the maxillae, there is a break in the bones where minerals go through the break. OM used this as a rough rotation point to shift the posterior halves of the maxillae back into place. Besides aiming for symmetry, it was hard to know how wide the jaw should be around the posteroventral processes of the maxillae. OM used the connection to the other bones (jugals, lacrimals, palatines, pterygoids, vomer) to figure out how wide it should be. Especially the palatines were very informative to know roughly how wide the skull should be at this point, since these needed to attach to the pterygoids medially and jugals, lacrimals and maxillae laterally.

- Bending the quadratojugals

There were always some bones that did not fit when OM put the skull together. In the end, OM rotated the dorsal process of the quadratojugals forward to the anterior position to make it fit with the squamosals and quadrates. The right quadratojugal was still in tight articulation with the jugal, so OM left that in place. The problem was that the angle of the dorsal process did not line up correctly with the quadrate and the squamosal. OM rotated this dorsal process around 10 degrees forward. The quadratojugal itself is a very thin element and did also seem to have sustained damage. OM felt fairly safe bending this bone instead of others that were less likely to have substantial changes to their shape during fossilization or after.

- Shifting of the right lacrimal and prefrontal

When viewed perfectly from the side, it becomes clear that the lacrimals did not line up. The right lacrimal is in good condition and showed little deformation. The left lacrimal was in much worse shape. OM opted to change the left lacrimal instead of the right one. The posterior margin of the left lacrimal is badly damaged. But the anterior margin is still in good shape and OM used that to align the two lacrimals. There was a gap, filled with matrix behind the prefrontal between it and the frontal bone. Since we know the nasals were bent to the right, it seems that it had taken the left lacrimal and tightly articulated prefrontal with it, pulling them forward. Thus, together with the nasals that needed to be bent to the left, OM needed to shift the left lacrimal and prefrontal back to a more posterior position that matched the right lacrimal and better articulated with the frontal.

Fixing small broken pieces of bone and putting them back into place to match the rest of the skull anatomy. Small pieces of bone were sometimes bent, or disarticulated entirely that needed to be changed.

Bones that OM bent

- The small remnant of the dentary was bent badly, OM straightened this.

- The ventral halves of the lacrimals were slightly bent due to larger stresses in the whole skull. OM laterally shifted these slightly so they articulated with the jugals and other bones better.

- The left jugal was in bad shape and OM laterally bent both the anterior half, and the posterodorsal process to better articulate with other bones.

- OM bent the right parasphenoid process to the right so it is oriented better. But OM left the smaller details as they were, as they don't articulate with anything else.

- OM did some very slight bending of the pterygoids to make them fit all the other bones they interact with (basisphenoid, ectopterygoid, palatine and vomer). OM felt okay with bending these slightly due to these bones being very thin and broken. It seems probable there was some deformation.

- OM very slightly bent the nasal and frontal dorsally around the point of the articulation with the nasal and prefrontals. Two reasons: There are several very small longitudinal cracks in the nasal that can be seen in the CT scans. This resulted in the whole complex of the braincase to move ventrally very slightly. As previously mentioned, the quadratojugal did not fit. Bending the nasal and frontal resulted in very slightly raising the whole braincase complex + postorbitals, squamosals and quadrates. This in turn made the quadratojugals articulate better and the posterodorsal process of the jugal articulates better with the ventral process of the postorbital.

Broken bones OM put back together

- OM used the breaks in the pterygoids to fit them around the rest of the bones. This resulted in a slightly smaller gap between the preserved parts.

- Small parts of the left postorbital and left squamosal were disarticulated. OM put these back as good as possible.

- Large parts of the vomer are missing. OM shifted the posterior piece back a little more so that it articulates better with the pterygoids. This results in a longer vomer.

- A small part of the ventromedial lip of the left maxilla was broken off. Initially, OM interpreted this as part of the vomer, but it seems to fit the maxilla better.

- Both ectopterygoids were broken and had a slight gap in the middle. OM narrowed those gaps to better reflect their original width.

SUPPLEMENTAL DATA 2

Characters for the phylogenetic analysis, modified from Rauhut and Pol 2019.

1. Premaxilla, height/length ratio ventral to external naris (length measured along ventral border of bone): < 0.5 (0), 0.5-0.99 (1), 1-1.49 (2), 1.5 or higher (3). Ordered.

2. Ventral margin of the premaxilla in lateral view: straight to slightly convex (0), markedly concave (1) (new).

3. Participation of the maxilla in the ventral border of naris: absent, subnarial process of the premaxilla contacts the nasal (0), subnarial process of premaxilla reduced or absent, maxilla expressed on the ventral margin of the nares (1) (modified from Rauhut and Pol 2019).

4. Premaxilla, posterior extent of nasal process relative to posterior tip of subnarial process: considerably more anterior (0), even (1), posterior (2). Ordered.

5. Premaxilla, form of premaxilla-nasal suture: V-shaped (0), W-shaped (1).

6. Position of anterior end of external nares: over anterior half of ventral margin of premaxillary body (0), over the posterior half of the ventral margin of the premaxillary body (1), posterior to the ventral margin of the premaxillary body (2). ordered.

7. Premaxilla, diastema adjacent to maxilla along dentigerous margin: absent (0), present, with alveolar margin being continuous between premaxilla and maxilla (1), present, with alveolar margin being discontinuous between premaxilla and maxilla ("subnarial gap") (2) (modified from Rauhut and Pol 2019).

8. Premaxilla, mediolateral constriction of posterior portion: absent (0), present (1).

9. Premaxilla, development of subnarial (maxillary) process: well-developed, rod or plate-like (0), reduced to a short, stout triangle, as long as or shorter than its basal width (1).

10. Subnarial foramen on the premaxilla-maxilla suture: absent (0), present (1).

11. Premaxilla, articulation with maxilla: planar (0), interlocking (1).

12. Anterior ramus of the maxilla: absent, anterior margin of maxillary body confluent with anterior margin of the ascending process (0), present but very short (length/height ratio less than 1) (1), present moderately long (1-1.35) (2), present, long (1.35-1.7) (3), present, very long (more than 2) (4). Ordered.

13. Subnarial fossa on the maxilla: absent (0), present (1) (new).

14. Maxilla, orientation of anteriormost alveolus: vertical (0), angled anteriorly (1).

15. Anterodorsal margin of the ascending process of the maxilla: straight or gently curved (0), with pronounced kink at about mid-length, with a more anteriorly facing margin ventral and a more dorsally facing margin dorsal to the kink (1).

16. Relative posterior length of the ascending process and jugal ramus of the maxilla: jugal ramus extends considerably further posteriorly than ascending process (0), subequal in length or ascending process longer (1) (new).

17. Maxilla, morphology of palatal process: long, ridged or fluted prong (0), long and plate-shaped (1).

18. Maxilla, position of palatal process: ventral, immediately dorsal to paradental plates (0), dorsal, immediately ventral to dorsal surface of maxillary anterior ramus (1).

19. Maxilla, horizontal ridge (prominent ‘lingual bar’) between palatal process and antorbital fenestra: absent (0), present (1).

20. Maxilla, depth of paradental plates relative to anteroposterior width: low, < 1.8 (0), tall > 1.8 (1).

21. Maxilla, ventral extent of paradental plates relative to lateral wall: as far ventral (0), fall short (1).

22. Maxilla, arrangement of nutrient foramina on lateral surface: single row or no distinct pattern (0), a second, more dorsally placed row is present anteriorly and converges with the ventral row posteriorly (1), a second, dorsally placed row is present and extends posteriorly more or less parallel to the ventral row (2).

23. Maxilla, anteroventral border of antorbital fossa: graded or stepped (0), demarcated by raised ridge (1).

24. Maxilla, anterior margin of antorbital fossa: rounded (0), squared (1).

25. Maxilla, ventral extent of antorbital fossa (as measured from the rim of the antorbital fenestra to the highest point of the ridge marking its border) at the level of about the half length of the antorbital fenestra: small or absent, less than 1/3 the height of the maxillary body at this level (0), moderate, between 1/3 and half the height (1), dorsoventrally deep, more than half the height (2). Ordered.

26. Maxilla, position of anterior end of antorbital fossa: posterior to or level with posterior rim of external nares (0), ventral to external nares (1).

27. Medial wall of the anterior end of the maxillary antorbital fossa: lacking depressions or foramina (0), with a large depression without sharply defined margins (1), with a sharply rimmed maxillary fenestra (2).

28. Development of maxillary fenestra: opens medially into a small maxillary antrum with a robust medial wall (0), opens medially into a large maxillary antrum that is medially open or only covered by a very thin bony wall (1), opens anteriorly into a large antrum within the ascending process of the maxilla (2) This character is inapplicable in taxa that lack a maxillary fenestra.

29. Promaxillary foramen: absent (0), present and opens anteriorly into pneumatic recesses in the ascending process of the maxilla (1).

30. Size of the promaxillary foramen in relation to the maxillary fenestra: smaller (0), larger (1). This character is inapplicable in taxa that lack a maxillary fenestra.

31. Maxilla, development of pneumatic fossa (excavatio pneumatica) in ascending process: absent (0), present (1).

32. Maxilla, pneumatic region on medial side of maxilla posteroventral to maxillary fenestra: absent (0), present (1).

33. Shape of the alveolar border of the maxilla: straight (0), convex (1), sinusoidal (2) (new).

34. Maxilla, posterior end of tooth row relative to orbit: beneath (0), anterior (1).

35. Posterior end of the jugal ramus of the maxilla: single (0), forked (1) (new).

36. Posteriormost end of the jugal ramus of the maxilla: straight, in line with alveolar border (0), markedly downturned (1) (new).

37. Maxilla and nasal, external surface texture: smooth (0), sculptured (1).

38. Nasal, inter-nasal contact in adults: separate (0), partly or fully fused (1).

39. Nasal, posterior narial margin: absent or weak fossa (0), large fossa (1), laterally splayed hood (2).

40. Posteriorly pointed, sharply rimmed depression on the lateral side of the nasal posterodorsal to the external nares: absent (0), present (1).

41. Nasal, participation in antorbital fossa: absent or at edge (0), present (1).

42. Nasal, antorbital fossa in lateral view: visible (0), occluded by ventrolaterally overhanging lamina (1).

43. Nasal, pneumatic foramina: absent (0), present (1).

44. Nasal, development of dorsolateral surfaces: none, nasals low and dorsally convex (0), pronounced dorsolateral rims, sometimes with lateral crests (1), tall, parasagittal crests (2) (modified from Rauhut and Pol 2019).

45. Midline crest on the nasals: absent (0), present (1) (modified from Rauhut and Pol 2019).

46. Posterior end of nasal crest on the nasal: not expanded, crest stays of subequal width throughout length (0), transversely broadened into a notable tubercle (1) (new).

47. Nasal, sculpturing: smooth or low rugosity (0), deeply rugose, bears large excresences (1) (modified from Rauhut and Pol 2019).

48. Posterior end of the nasals: thin, plate-like (0), strongly thickened, robust (1) (new).

49. Antorbital fossa and dorsal rim on the anterior process of the lacrimal: present, but fossa is largely hidden in lateral view by an overhanging lateral lamina and only exposed anteriorly (0), present, widely exposed laterally and confluent with the antorbital fossa of the dorsal part of the ventral process (1), dorsal rim absent, no differentiation between fossa and rim on the lateral surface of anterior process (2).

50. Lacrimal, morphology of lateral lamina of ventral process of lacrimal: forming a continuous sheet of bone between the ventral and anterior processes (0), invaginated dorsally and convex anteriorly, anteriormost point situated dorsal to midheight of ventral process (1), anteriormost point situated around midheight of ventral process (2).

51. Anteroposterior expansion of ventral process of the lacrimal: begins at two thirds of the height of the ventral process or lower (0), begins at the dorsal end of the ventral process (1) (new).

52. Lacrimal, dorsal and ventral portions of antorbital fossa: separated by anterior projection of lateral lamina (0), continuous, lateral lamina does not project far anteriorly (1).

53. Lacrimal fenestra: absent (0), present (1).

54. Lacrimal, openings in lacrimal recess: single (0), multiple (1).

55. Lacrimal horn: absent (0), small dorsal rugosity (1), low, broad, rugose bar (2), large triangular horn (3). Ordered.

56. Lacrimal, suborbital process: absent (0), present (1).

57. Lacrimal, angle between anterior and ventral rami: ~90° (0), < 75° (1).

58. Lacrimal, length of anterior process relative to ventral process: more (0), or less than 80% (1).

59. Jugal, position of anterior end: excluded from internal antorbital fenestra (0), posterior to internal antorbital fenestra, but reaching its posterior rim (1), expressed at rim of internal antorbital fenestra, with distinct anterior process extending beneath it (2) (modified from Rauhut and Pol 2019). Ordered.

60. Jugal, pneumatisation: absent (0), internally hollowed and transversely inflated by foramen in posterior rim of antorbital fossa (1).

61. Jugal, antorbital fossa that is clearly offset from the lateral jugal surface by a raised rim: absent (0), present (1) (modified from Rauhut and Pol 2019).

62. Anterior end of jugal: slender and not or only slightly expanded (0), strongly expanded, and expansion forms at least a small part of the anterior margin of the orbit (1).

63. Jugal, orientation of orbital margin on postorbital process: angled posterodorsally (0), vertical (1) (modified from Rauhut and Pol 2019).

64. Horizontal ridge along the jugal body: absent (0), present (1) (new).

65. Lateral ridge along the postorbital contact on the postorbital process of the jugal: absent (0), present (1) (new).

66. Dorsoventral height of the posterior process of jugal: less than half the height of the suborbital part (0), more than half the height, but less than height of suborbital part (1), or subequal or more than dorsoventral height of suborbital part (2) (modified from Rauhut and Pol 2019).

67. Postorbital, articulation with jugal: planar or with a shallow, V-shaped groove dorsally (0), with a deep, broad groove posteriorly, ventral process with U-shaped cross-section (1).

68. Postorbital, suborbital flange: absent (0), present (1).

69. Development of suborbital flange of postorbital: small, triangular eminence (0), dorsoventrally elongate, large rounded flange (1), jugal process curved anteroventrally and suborbital process developed as large, sharply angled, triangular flange (2). This character is not applicable to taxa that do not have a suborbital flange.

70. Postorbital, ventral extent relative to ventral margin of orbit: substantially above (0), approximately same level (1).

71. Dorsal margin of postorbital above jugal process in lateral view: convex (0), straight to slightly concave (1), with pointed, triangular cornual process that is anteriorly offset from a possible step at the beginning of the squamosal articulation (2) (new).

72. Laterosphenoid facet on medial side of the postorbital: large and placed at the junction of the three postorbital processes (0), reduced in size and placed entirely on the anterior process (1) (new).

73. Medial side of posterior process of the postorbital: straight or concave, dorsal margin forms a sharp rim (0), convex, medial margin curves into dorsal margin (supratemporal fossa extends onto process) (1).

74. Supratemporal fossa on the anterior process of the postorbital and posterior end of frontal: forms a large shelf on the dorsal surface of the process (0), reduced, restricted to the posteriormost part and faces more posterodorsally than dorsally (1).

75. Supraorbital brow: absent, anterior end of postorbital tapers (0), present as a dorsoventrally expanded, anteriorly rounded, rugose swelling over the posterior part of the orbit (1), large, strongly rugose supraorbital brow (possible formed by a separate palprebal ossification fused to the postorbital) that connects the postorbital with the lacrimal present (2). Ordered.

76. Anterior process of the postorbital: straight (0), downturned anteriorly (1) (new).

77. Posterior process of the postorbital: Pointed and transversely narrow (0), broadened transversely posteriorly and wider than high (1).

78. Parietal-postorbital contact at the anterior end of the supratemporal fenestra: absent, parietal and postorbital separated by posterior process of frontal that reaches the laterosphenoid (0), present, frontal excluded from laterosphenoid in dorsal view (1).

79. Prefrontal in adult individuals: Exposed on the anterodorsal margin of the orbit (0), reduced, not exposed at the anterior margin of the orbit, might only be visible at the dorsal margin of the orbit (1), absent as separate ossification (2).

80. Frontal-prefrontal contact: prefrontal contacts anterolateral margin of the frontal, placed mainly lateral to the frontal (0), prefrontal contacts anterior margin of the frontal, placed mainly anterior to frontal (1) (new).

81. Frontal, exposure along orbital rim: broad, one third or more of the dorsal orbital margin (0), reduced, less than one third of the dorsal orbital margin (1).

82. Midline ridge on the frontal: absent (0), present (1) (new).

83. Proportions of frontal (as exposed on the skull roof): longer than wide (0), wider than long (1).

84. Frontals in adult individuals: unfused (0), partially or completely fused (1) (Carrano and Sampson 2008).

85. Clearly offset lateral postorbital process of the frontal: absent (0), present (1) (new).

86. Nasal-frontal suture: anterior end of articulated frontals medially tapering or rounded (0), anterior end of articulated frontals separated by a broad medial posterior process of the nasal (1).

87. Overlap of the parietal over the median supraoccipital ridge/thickening: absent (0), present (1) (modified from Rauhut and Pol 2019).

88. Parietal, development of median skull table between supratemporal fossae: flat, but relatively narrow (less than 50% of width of supratemporal fenestrae) (0), separated by a triangular plate of bone anteriorly, but narrowing to a sagittal crest posteriorly (1), narrow with sharp sagittal crest (2), very broad, widely separating upper temporal fenestrae, skull table at least 50% of width of supratemporal fenestra and fossa (3).

89. Parietal, size and elevation of nuchal wedge and alae: small to moderate, height of alae less than height of base of paroccipital process (0), expanded, height more than height of base of paroccipital process (1).

90. Supratemporal fossa, anteromedial corner: open dorsally (0), partially roofed over by a small shelf of the frontalparietal (1).

91. Squamosal, constriction of lower temporal fenestra: absent (0), present (1).

92. Squamosal, anterodorsal lamina: emarginated by upper temporal fenestra (0), continuous (1).

93. Strongly developed and sharply defined horizontal ridge extending from the ventral margin of the postorbital facet towards the quadrate articulation on the lateral side of the squamosal body: absent (0), present (1) (new).

94. Posterior extent of postorbital facet on squamosal: ends anterior to or at the anterior border of the ventral process (0), extends posteriorly to at least the half-width of the ventral process (1) (new).

95. Relative length of anterior and posterior processes of squamosal: anterior process considerably longer than posterior process (0), anterior process as long as or shorter than posterior process (1) (new).

96. Squamosal, flange covering quadrate head laterally: absent (0), present, covers the posterior part of the head and separated from quadratojugal process of squamosal by a wide, U-shaped incision (1), present, covers most of quadrate head and separated from quadratojugal process only by a narrow, slit-like incision (2). Ordered.

97. Squamosal-quadratojugal contact: present (0), absent (1) (modified from Rauhut and Pol 2019).

98. Squamosal, articulation with quadratojugal: at tip (0), broad (1) (modified from Rauhut and Pol 2019).

99. Quadratojugal, anteriormost point of ventral process relative to lower temporal fenestra: ventral, no further than two thirds of the length of the fenestra from the posterior border (0), further anterior, subequal to the anterior border of the fenestra (1).

100. Large pneumatic foramen in quadrate: absent (0), present (1). (modified from Rauhut and Pol 2019).

101. Position of the quadrate head in relation to the orbit: low, below two-thirds of the height of the orbit (0), at two-thirds of the height of the orbit or higher (1).

102. Angle between quadrate ridge and distal quadrate condyles in posterior view: more than 70° (0), less than 70° (1) (modified from Rauhut and Pol 2019).

103. Medial side of quadrate ridge at mid-height in posterior view: straight or concave (0), convex (1) (new).

104. Proportion of the length of the quadrate that is occupied by the pterygoid wing: less than 70% (0) or 70% or more (1).

105. Proximal part of the dorsal margin of the pterygoid wing of the quadrate: forms a sharp crest (0), notably thickened mediolaterally (1) (new).

106. Ventral margin of pterygoid wing of the quadrate: Offset from mandibular condyles (0), confluent with expansion for mandibular condyles (1) (new).

107. Ventral margin of the pterygoid wing of the quadrate: flexed medially to form a medial shelf or ridge (0), forms a sharp ventral edge , only the most proximal part flexes medially (1) (new).

108. Quadrate foramen: present (0), absent(1) (modified from Rauhut and Pol 2019).

109. Development of quadrate foramen in the quadratojugal flange of the quadrate: semioval notch, widely open laterally (0), with a ventrally flexed lateral spur at at least the dorsal margin (1), completely enclosed in the quadrate (2) (modified from Rauhut and Pol 2019).

110. Orientation of the quadrate in lateral view: anteroventrally inclined, mandibular joint notably anterior to quadrate head (0), more or less vertical, mandibular joint approximately straight below quadrate head (1), posteroventrally inclined, mandibular joint notably posterior to quadrate head (2) (modified from Rauhut and Pol 2019).

111. Quadrate, head shape in dorsal view: oval (0), subrectangular (1).

112. Mediolateral expansion of quadrate head in relation to quadrate shaft: absent (0), present (1) (new).

113. Quadrate, medial foramina adjacent to condyles: absent (0), present (1).

114. Medial condyle of the quadrate: anteroposteriorly narrow and long axis of condyle mainly mediolaterally oriented (0), anteroposteriorly expanded and more anteroposteriorly oriented (1) (new).

115. Ventral rim of the basis of the paroccipital processes: above or level with the dorsal border of the occipital condyle (0), situated at mid-height of occipital condyle or lower (1).

116. Paroccipital processes: directed laterally or slightly ventrolaterally (0), directed strongly ventrolaterally, with distal end entirely below the level of the foramen magnum (1).

117. Marked depression on the exoccipital lateral to the foramen magnum, above the paracondylar recess: present (0), absent (1) (new).

118. Supraoccipital, anteroposterior depth of median ridge relative to occipital condyle length: less (0), greater (1).

119. Maximal width of dorsal expansion of supraoccipital ridge: less than or subequal to width of foramen magnum (0), notably greater than width of foramen magnum (1).

120. Posterior exit of mid-cerebral vein: on the posterior surface of the supraoccipital, without any associated marked depressions (0), placed within a notable, funnel-shaped depression (1), associated with a curved groove leading towards the posttemporal fenestra (2) (new).

121. Supraoccipital, participation in foramen magnum: present, ventral margin of supraoccipital forms a more or less straight line above the foramen, forming most of its dorsal rim (0), narrow, formed by a small median ventral process of the supraoccipital separating the exoccipitals on the dorsal edge of foramen (1), absent, exoccipitals meet in the midline above the foramen magnum (2) (modified from Rauhut and Pol 2019). ordered.

122. Attachment for proatlas on dorsolateral rim of foramen magnum: inconspicuous (0), marked as paired raised lips (1) (new).

123. Basioccipital, ventrolateral pair of pneumatic cavities invading neck of occipital condyle and joining medially: absent (0), present (1).

124. Morphology of posterior basioccipital surface below the condyle: with undivided longitudinal median groove (0), median groove divided dorsally by small median lamina (1), with large longitudinal ridge separating two large lateral depressions (2).

125. Basioccipital, fossa ventral to occipital condyle in basioccipital apron: narrow and groove-like, one half or less of the width of the occipital condyle (0), broad depression, approximately two thirds or more the width of the occipital condyle (1) (Carrano et al. 2012).

126. Notch on the basioccipital-basisphenoid suture separating a medial basioccipital portion of the basal tubera from a lateral basisphenoid portion: absent (0), present (1) (modified from Rauhut and Pol 2019).

127. Width of basioccipital ventral to the occipital condyle: greater than width of occipital condyle (0), subequal to or less than the width of the occipital condyle (1).

128. Basisphenoid, location of basipterygoid processes relative to basal tubera: anterior or slightly anteroventral, basisphenoid recess opens ventrally (0), anteroventrally, basisphenoid recess opens posteroventrally (1), almost directly ventral, basisphenoid recess anteroposteriorly narrower than wide and opens more posteriorly than ventrally (2).

129. Basisphenoid, presence and depth of basisphenoid recess: absent (0), shallow, longer than deep (1), deep, deeper than long (2). Ordered.

130. Basisphenoid, shape of opening for basisphenoid recess: elongate oval (0), teardrop-shaped, narrowing posteriorly (1), subcircular (2), anteroposteriorly compressed, slit-like (3), trapezoidal, widening posteriorly (4).

131. Longitudinal lamina at least partially dividing the basisphenoid recess in its interior into left and right compartments: present (0), absent (1) (new).

132. Subdivision of basisphenoid recess into anterior and posterior recesses: absent (0), present (1) (new).

133. Posterior part of the ventral margin of the basisphenoid between the basal tubera and the basipterygoid processes: gently concave in lateral view (0), straight or slightly convex in lateral view (1).

134. Basipterygoid processes: on elongate stalks, with rather small articular surface (0), broad, enlarged articular surface facing anteroventrolaterally at lateral sides of the anterior end of the basisphenoid (1) (modified from Rauhut and Pol 2019).

135. Exit of cranial nerves X and XI: laterally through the metotic foramen (0), posteriorly through a foramen in the paracondylar recess (1).

136. Subcondylar recess on the occiput: small, restricted to the occiput lateral to the occipital condyle (0), extended ventrally, developed as deep depression on the lateral sides of the ventral part of the occiput (1)

137. Orientation of the occipital condyle in respect to the skull table: posteriorly (0), posteroventrally (1) (modified from Rauhut and Pol 2019).

138. Angle between the posterior end of the dorsal skull roof and the main body of the supraoccipital: approximately 90° (0), notably obtuse (1).

139. Braincase, morphology of trigeminal foramen: single (0), partly split (1), fully split (2). Ordered.

140. Exit of the abducens nerve (cranial nerve VI): placed within the pituitary fossa or the depression surrounding it (0), placed lateral to the pituitary fossa and its surrounding depression (1).

141. Dorsal process on the parasphenoid rostrum anterior to the pituitary fossa: absent (0), present (1) (new).

142. Well-developed anterior tympanic recess in the braincase: absent (0), present (1).

143. Braincase, ossification of interorbital region: weak or absent (0), extensive, ossified sphenethmoid and interorbital septum (1).

144. Length of the anterior, maxillary process of the palatine (as measured from the anterior end of the junction with the vomerine process to the anterior tip) in relation to length of jugal process (as measured from the posterior end of the junction with the pterygoid process to the posterio tip): less or subequal (0), longer (1).

145. Palatine, pneumatic recess on the dorsolateral side of the base of the vomerine process: absent (0), present (1) (modified from Rauhut and Pol 2019).

146. Deep, posterodorsally opening recess at the confluence of the jugal, pterygoid and vomerine processes of the palatine: absent (0), present (1) (new).

147. Pterygoid, pocket on ectopterygoid flange: absent (0), present (1).

148. Anteroventral expansion of jugal process of ectopterygoid: absent (0), present (1)

149. Ectopterygoid, ventral fossa: absent (0), present (1).

150. Ventral fossa of the ectopterygoid: simple depression (0), invaginates the lateral ectopterygoid body (1).

151. Size of external mandibular fenestra: Large, surangular above the fenestra accounts for less than half of the height of the mandible (0), reduced, surangular accounts for more than half the height of the mandible (1).

152. Mandible, position of anterior end of external mandibular fenestra relative to last dentary tooth: posterior (0), ventral (1).

153. Anterior end of dentary: not expanded (0), slightly expanded (1), strongly dorsally expanded, dorsal margin considerably above posterior alveolar margin (2) (modified from Rauhut and Pol 2019). Ordered.

154. Shape of the anterior end of the dentary: rounded (dorsoventrally convex) (0), squared (dorsoventrally straight) (1).

155. Anteroventral flange or process at the anterior end of the dentary: absent (0), present (1).

156. Enlarged tooth or teeth in the anterior end of the dentary: absent (0), present, usually in the third and/or fourth dentary alevolus (1).

157. Dentary, shape in dorsal view: straight (0), curves anteromedially (1).

158. Dentary, longitudinal groove housing dorsally situated row of neurovascular foramina on lateral surface: absent or weak (0), present and well-defined (1).

159. Dentary, number of Meckelian foramina: one (0), two (1).

160. Posterior end of the dentary: with dorsal and ventral processes subequal in length or dorsal process slightly longer (0), sloping posteroventrally, ventral end extends considerably further posteriorly than dorsal end (1).

161. Morphology of posterior margin of dentary: forked (0), posteroventrally sloping margin with incision for mandibular fenestra (1).

162. Splenial, contour of posterior edge: straight (0), curved (1), notched (2).

163. Splenial, size of splenial (‘mylohyoid’) foramen: small (height less than 15 % of height of splenial at the level of the foramen) (0), intermediate (height between 15 % and 25 % (1), large (height 25% or more) (2). Ordered.

164. Splenial, foramen in ventral part: completely enclosed by bone (0), open anteroventrally (1).

165. Deep incision in the anteroventral part of the prearticular: absent (0), present (1) (new).

166. Surangular, horizontal ridge on lateral surface below mandibular joint: weak or absent (0), strong (1).

167. Surangular, depressed lateral shelf for the attachment of the M. adductor mandibulae externus superficialis, bound medially by a dorsally facing ridge: absent, dorsal surface of the surangular mediolaterally convex anterior to the glenoid (0), present (1) (new).

168. Shelf on surangular for the attachment of the M. adductor mandibulae externus superficialis: faces mainly dorsally (0), faces notably laterodorsally (1)

169. Surangular, number of posterior surangular foramina: one (0), two (1).

170. Articular: rectangular ventral projection below the foramen for the chorda tympani on the medial side at the posteriomedial margin of the mandibular glenoid: absent, ventromedial margin of glenoid rounded (0), present (1) (modified from Rauhut and Pol 2019).

171. Anterior rim of the mandibular glenoid in lateral view: confluent with the dorsal margin of the surangular (0), raised above the dorsal margin of the surangular (1).

172. Ossified antarticular in the mandible: absent (0), present (1).

173. Retroarticular process of the mandible: elongate, as long as or longer than anteroposterior length of mandibular glenoid (0), short, shorter than length of mandibular glenoid (1).

174. Retroarticular process, mediolateral width relative to posterior width of dentary: ≤ (0), > (1).

175. Retroarticular process, orientation of attachment surface: posterodorsal (0), posterior (1).

176. Attachment area for the M. depressor mandibulae: mediolaterally notably concave (0), flat to slightly convex (1).

177. Lateral margin of retroarticular process: more or less aligned with glenoid (0), strongly offset ventrally from glenoid (1).

178. Paradental plates, continuity and replacement groove: separated, groove present (0), forming a continuous medial lamina (‘fused’), groove absent (1).

179. Interdental plates, visibility in medial view: widely exposed, subpentagonal and moderate-tall (0), obscured by an expanded paradental lamina, triangular apices only may be visible (1).

180. Paradental plates, surface texture: smooth (0), vertically striated or ridged (1).

181. Lateral maxillary and dentary teeth: recurved, so that tip of the tooth is placed distal to distal carina, distal carina concave (0), straight or almost straight, tip of tooth placed mesial to distal carina, distal carina straight or convex (1).

182. Teeth, crown striations: absent (0), present on premaxillary and/or anterior dentary teeth only (1), present on all tooth crowns (2).

183. Teeth, enamel wrinkles: absent (0), pronounced marginal enamel wrinkles (1).

184. Lateral teeth, ratio between mesiodistal length and transverse width at the base: more than 1.5 (0), 1.5 or less (1) (modified from Rauhut and Pol 2019).

185. Teeth, maxillary and dentary, serrations: present (0), absent (1).

186. Teeth, maxillary and dentary, extent of anterior carina: to base of crown (0), at mid-height of crown or more apically (1).

187. Premaxillary teeth, arrangement of carinae: nearly symmetrical, on opposite sides (0), more asymmetrical, both on lingual side (1).

188. Premaxillary teeth, serrations: present (0), absent (1).

189. Premaxillary teeth, number: three (0), four (1), five (2), more than five (3). Ordered.

190. Premaxillary teeth, spacing: even (0), paired and spaced (1).

191. Size of first premaxillary tooth (or alveoli): subequal to second premaxillary tooth (0), less than two thirds of the size of the second premaxillary tooth (1), less than half the size of the second premaxillary tooth (2). Ordered.

192. Maxillary teeth, number: > 17 (0), 11-17 (1), < 11 (2). Ordered.

193. Maxillary teeth, mid-tooth spacing: adjacent, closely spaced (0), more widely spaced in the mid-section (at least one-third of an alveolar width apart), but spacing decreases posteriorly again (1), more widely spaced, spacing increases posteriorly towards the end of the tooth row (2) ordered.

194. Rapid increase in tooth size at the anterior end of the maxilla: absent (0), present (1) (new).

195. Dentary teeth, size and number relative to maxillary teeth: approximately equal (0), smaller and approximately 1.5 times as numerous (1).

196. Cervical vertebrae, anterior articular facet: concave (0), flat (1), convex (2). Ordered.

197. Pneumatic feature posterior or posterodorsal to parapophysis (anterior pleurocoel) in cervical vertebrae: absent (0), large, blind depressions (1), large foramina (2).

198. Pneumatic feature on the posterior half of the vertebral centrum in cervical vertebrae: absent (0), large depression (1), foramen (2).

199. Presacral vertebrae, extent of anterior pleurocoel: anterior dorsals only (0), to sacrum (1).

200. Vertebrae, internal structure of pneumatic centra: absent (0), camerate (1), camellate (2).

201. Epipophysis of the atlantal neural arch in lateral view: slender, rod-like (0), expanded, triangular (1).

202. Axial neural spine: anteroposteriorly extensive, sheet-like, with convex or only gently ascending dorsal margin (0), anteroposteriorly reduced, rod-like, with steeply ascending dorsal margin (1).

203. Axis, orientation of intercentrum ventral surface: horizontal or slightly anteroventral (0), tilted anterodorsally (1).

204. Length of epipophyses of the axis: short, approximately level with posterior end of postzygapophyses (0), long, overhang postzygapophyses posteriorly for more than half the length of the postzygapophyseal articular facet (1).

205. Spinopostzygapophyseal laminae of the axis: Extensive, connecting the spine with the postzygapophyses in a large arch, resulting in a large, triangular to rhomboid fossa on the posterior side of the neural arch (0), reduced, strongly invaginated between spine and postzygapophyses, so that at least the distal part of the spine has parallel borders and the posterior fossa is reduced (1).

206. Development of parapophysis on axis: well-developed facet on the anteroventral side (0), indistinct, or only developed as a slightly roughened patch (1).

207. Development of diapophysis on axis: indistinct, probably absent (0), developed as distinct ventrolateral projection (1).

208. Axis, pleurocoels: absent (0), present (1).

209. Neural spine of third cervical vertebra: not significantly different from other postaxial cervical neural spines (0), slender and strongly backswept (1)

210. Cervical vertebrae, morphology of anterior pleurocoel: single opening (0), two openings oriented anteroventralposterodorsal or very plastic morphology (1).

211. Cervical vertebrae, middle, shape of anterior pleurocoel: round (0), anteroposteriorly elongate (1).

212. Cervical vertebrae, anterior, ventral keel: present (0), absent or weak ridge (1).

213. Cervical vertebrae, anterior, demarcation of dorsal surface of neural arch from diapophyseal surface: gently sloping (0), pronounced edge or ridge that offsets the dorsal from the lateral surface (1), elevated prezygoepipophyseal lamina, dorsal surface concave between neural spine and the lamina (2). Ordered.

214. Cervical vertebrae, articular surface of prezygapophyses: planar (0), flexed (1).

215. Cervical vertebrae, perimeter of anterior articular surface: not rimmed by a flattened peripheral band (0), flat, forming a distinct rim (1).

216. Cervical vertebrae, anterior, transverse distance between prezygapophyses relative to width of neural canal: < (0), >, prezygapophyses situated lateral to neural canal (1).

217. Cervical vertebrae, anterior, morphology of epipophyses: low, wider than high, posteriorly pointed (0), transversely narrow, high (1), high, robust (2).

218. Neural spine of mid-cervical vertebrae: anteroposteriorly longer than dorsoventrally high (0), higher than long (1).

219. Cervical vertebrae, longest post-axial elements: first five (0), last five (1).

220. Length/posterior height ratio of mid-cervical centra: less than 1.75 (0), 1.75-2.75 (1), more than 2.75 (2). Ordered.

221. Width/height ratio of anterior articular surface of posteriormost cervical and anteriormost dorsal vertebrae: less than 1.3 (0), more than 1.3 (1) (new).

222. Height of anterior dorsal neural spines (as measured from the dorsal margin of the postzygapophysis): less (0) or more (1) than 1.25 times the height of the neural arch (as measured from the dorsal rim of the centrum to the dorsal margin of the postzygapophysis).

223. Ventral keel in posterior-most cervicals and anterior-most dorsals: ventrally concave, a rounded hypapophysis might be present anteriorly (0), forming a straight to slightly convex ventral margin, anterior end of keel protrudes ventrally from the anterior articular surface and is separated from the latter by a distinct step (1).

224. Dorsal vertebrae, pneumaticity/webbing at base of neural spines in middle to posterior dorsals: absent (0), present (1).

225. Middle to posterior dorsal vertebrae, accessory centrodiapophyseal lamina: absent (0), present (1).

226. Dorsal vertebrae, anterior, ventral keel: absent or developed as a weak ridge (0), pronounced, around 1/3 the height of centrum and inset from lateral surfaces (1).

227. Dorsal vertebrae, anterior, size of pneumatic foramen in centrum: small (0), enlarged (1).

228. Dorsal vertebrae, elevation of parapophyses: slightly elevated from centrum (0), project far laterally, more than half the diapophyseal length (1).

229. Dorsal vertebrae, orientation of hyposphene laminae: diverge ventrolaterally (0), parallel and sheet-like (1).

230. Dorsal vertebrae, position of parapophyses in posteriormost elements: on the same level as transverse process (0), distinctly below transverse process (1).

231. Dorsal vertebrae, distinct step-like ridge lateral to hyposphene, running posterodorsally from dorsal border of neural canal to posterior edge of postzygapophyses: absent (0), present (1), ridge present and is developed into a prominent lamina that bisects the infrapostzygapophyseal fossa in posterior dorsal vertebrae (2). Ordered.

232. Posterior dorsal vertebrae, shape of ventral surface of vertebral centra: transversely rounded (0), flattened, sometimes with a shallow medial sulcus (1) (new).

233. Dorsal vertebrae, morphology of neural spines: transversely compressed sheets (0), transversely broad anteriorly and posteriorly, central regions of lateral surface embayed by deep vertical troughs (1).

234. Dorsal vertebrae, posterior, inclination of neural spines: vertical or posterior (0), anterior (1).

235. Dorsal vertebrae, height of neural spines relative to centrum height: low, ≤ 1.3x (0), moderate, 1.4-1.8x (1), tall, ≥ 2.0x (2). Ordered.

236. Middle to posterior dorsal vertebrae, centrum length relative to height: centrum higher than long (0), centrum longer than high (1), centrum more than 1.5 times longer than high (2) (modified from Rauhut and Pol 2019). ORDERED.

237. Sacral vertebrae, centrum pneumaticity: absent (0), pleurocoelous fossae (1), pneumatic foramina (2).

238. Sacral vertebrae, number: 2 [primordial sacrals only] (0), 5 [1 dorsosacral, 2 caudosacrals] (1), 6 [2 dorsosacrals, 2 caudosacrals] (2). Ordered.

239. Sacral vertebrae, transverse dimensions of middle centra relative to other sacrals: equivalent (0), constricted (1).

240. Sacral vertebrae, orientation of ventral margin of middle centra: approximately horizontal (0), strongly arched (1).

241. Sacral vertebrae, dorsal edge of neural spines: as thin as remainder of spine (0), transversely thickened (1).

242. Sacral vertebrae, pneumaticity of neural arches: weak or absent (0), paired fossa ventral to diapophyses (1).

243. Caudal vertebrae, anterior, morphology of ventral surface: flat (0), groove (1), ridge (2).

244. Caudal vertebrae, L-shaped neural spines: absent (0), present (1).

245. Anterior to mid-caudal vertebrae, depressions or pneumatic foramina in centrum: absent (0), large, pronounced pleurocentral depressions on the dorsal part of the lateral side (1), pneumatic foramina (2).

246. Caudal vertebrae, anterior, centrodiapophyseal laminae on neural arch: weak or lacking (0), only anterior centrodiapophyseal lamina well developed, defining a deep prezygodiapophyseal fossa (1), as prominent as in dorsal vertebrae, defining deep infradiapophyseal fossa that penetrates neural arch (pneumatic) (2).

247. Spinodiapophyseal ridge or lamina in at least mid-caudal vertebrae: absent (0), present (1) (new).

248. Caudal vertebrae, anterior, proportions of neural arch base relative to centrum proportions: < (0), ≥ (1).

249. Caudal vertebrae, middle, morphology of neural spines: rod-like and posteriorly inclined (0), subrectangular and sheet-like (1), rod-like and vertical (2).

250. Cervical ribs, length of anterior process: short (0), long (1).

251. Gastralia, posteriormost gastral segments: separate (0), united into single, boomerang-shaped elements (1).

252. Sacral ribs, articulations in adults: separate (0), fused together (1).

253. Sacral ribs, position of posterior attachment to ilium: ventral (0), posterodorsal (1).

254. Sacral ribs, depth relative to ilium height: < 85% (0), ≥ 90% (1).

255. Chevrons, morphology in middle caudal vertebrae: rodlike or only slightly expanded ventrally (0), L-shaped (1).

256. Chevrons, proximal articular surface: divided into anterior and posterior facets by distinct transverse ridge (0), no ridge, but low lateral mounds may be present, one on each side (1).

257. Chevrons, curvature: straight or gently curved (0), strongly curved (1).

258. Chevrons, anterior process: absent (0), present (1).

259. Chevrons, morphology of distal end in anterior and middle elements: expanded anteroposteriorly (0), unexpanded, tapers ventrally (1).

260. Scapula, angle between blade and acromion: gradual, oblique (0), abrupt, perpendicular (1).

261. Scapula, size of acromion process: moderate (0), marked (1).

262. Scapula, midshaft expansion of blade: absent (0), present (1).

263. Scapula, distal expansion of blade: marked (0), weak/absent (1).

264. Scapula, length:width ratio of blade: ≤ 7 (0), 7.5-9 (1), > 10 (2). Ordered.

265. Scapulocoracoid, shape of anterior margin: indented or notched between acromial process and coracoid suture (0), smoothly curved and uninterrupted across scapula-coracoid contact (1).

266. Scapulocoracoid, glenoid lip: moderate (0), marked (1).

267. Coracoid, development of posteroventral process: low, rounded posteroventral eminence (0), pronounced, posteroventrally tapering process (1).

268. Coracoid, development of biceps tubercle (= acrocoracoid process): absent or poorly developed (0), conspicuous and well developed as tuber (1), developed as a posteroventrally oriented ridge (2).

269. Coracoid, prominent fossa on ventral surface posteroventral to glenoid (subglenoid fossa): absent (0), present (1).

270. Humerus, shape of head: elongate (0), globular (1).

271. Humerus, longitudinal torsion of shaft: absent (0), present (1).

272. Humerus, size of trochanters relative to midshaft diameter: < (0), > 150% (1) > 250% (2). Ordered.

273. Humerus, development of internal tuberosity: low/rounded (0), hypertrophied (1).

274. Humerus, length of deltopectoral crest relative to total bone length: < 0.4 (0), 0.43-0.49 (1) > 0.52 (2). Ordered.

275. Humerus, development of deltopectoral crest: large rectangular crest (0), reduced to a low, rounded flange (1).

276. Humerus, orientation of deltopectoral crest apex: anteriorly (0), anterolaterally (1).

277. Humerus, relative orientation of proximal and distal condyles in anteroposterior view: parallel, humerus straight (0), distal canted (1).

278. Humerus, anterior surface of bone adjacent to ulnar condyle: smooth or gently depressed (0), bears well-defined fossa (1).

279. Humerus, shape of distal condyles: rounded (0), flattened (1).

280. Radius and ulna, development of radial external tuberosity and ulnar internal tuberosity: low, rounded (0), hypertrophied distal ends of radius and ulna broadened (1).

281. Radius, shaft: straight (0), curves laterally (1).

282. Radius, development of medial biceps tubercle: small or indistinct (0), hypertrophied (1).

283. Ulna, olecranon process: absent (0), present (1).

284. Ulna, morphology of olecranon process: transversely robust (0), transversely compressed and ‘blade-like’ (1).

285. Ulna, crest extending distally along posterior surface from olecranon process: absent (0), present (1).

286. Ulna, hypertrophied medial and lateral processes on proximal end: absent (0), present (1).

287. Ulna, length relative to minimum circumference: stout, < 2.3 (0), gracile > 2.6 (1).

288. Carpus, morphology and articulations of distal carpals: separate dc1 and dc2 over separate metacarpals, flattened proximodistally (0), fused dc1 and dc2, dc1 overlaps metacarpals I and II, flattened proximodistally (1), fused dc1 and dc2, dc1 overlaps metacarpals I and II, strongly arched proximodistally (2).

289. Manus, length of digit II relative to length of humerus: < (0), > (1)

290. Manus, composition: digit IV and V present (0), digit IV present, digit V absent (1), MC IV present, IV phalanges and digit V absent (2), digits IV and V absent (3). Ordered.

291. Manual digits, lengths: III longest (0), II longest (1).

292. Metacarpals, transverse width of proximal articular ends relative to minimum transverse shaft width: < (0), ≥ 2x (1).

293. Metacarpal I, length to minimum width ratio: 1.4-1.9 (0), ≥ 2.4 (1).

294. Metacarpal I, length relative to length of metacarpal II: > 57% (0), < 57% (1).

295. Metacarpal I, extent of contact with metacarpal II relative to shaft length: < 1/3 (0), 1/2 (1).

296. Metacarpal I, angle between facet for metacarpal II and proximal articular facet: perpendicular (0), obtuse (1).

297. Metacarpal III, position of base relative to those of other metacarpals: at same level (0), on palmar surface (1).

298. Metacarpal III, shape of proximal end: rectangular (0), triangular (1).

299. Metacarpal III, width relative to width of metacarpal II: > 50% (0), < 50% (1).

300. Manual ungual I, length:height ratio: < 2.5x (0), > 2.5x (1).

301. Manual unguals, proximal height:width ratio: transversely broad, < 2.0 (0), transversely narrow, > 2.4 (1).

302. Pelvic elements, articulations in adults: separate (0), fused (1).

303. Ilium, large external pneumatic foramina and internal spaces: absent (0), present (1).

304. Ilium, vertical ridge on lateral surface of blade dorsal to acetabulum: absent (0), low swollen ridge (1), low double ridge (2).

305. Ilium, posterior width of brevis fossa: subequal to anterior width, fossa margins subparallel (0), twice anterior width, fossa widens posteriorly (1).

306. Ilium, height of lateral wall of brevis fossa relative to medial wall: taller along whole length (0), shorter anteriorly, exposing medial wall in lateral view (1).

307. Ilium, morphology between supraacetabular crest and brevis shelf on lateral surface: gap (0), continuous ridge (1).

308. Ilium, ventrolateral development of supraacetabular crest: large/pendant ‘hood’ (0), reduced shelf (1).

309. Ilium, orientation of pubic peduncle: mostly ventral (0), mostly anterior or ‘kinked’ double facet with anterior and ventral components (1).

310. Ilium, shape of acetabular margin of pubic peduncle: transversely convex or flat (0), transversely concave (1).

311. Ilium, relative sizes of pubic and ischial articulations: subequal (0), pubic articulation ≥ 130% of iliac articulation (1).

312. Ilium, morphology of ischial peduncle: rounded (0), acuminate (1).

313. Ilium, pubic peduncle length to width ratio: ≤ 1 (0), 1.3-1.75 (1), > 2 (2). Ordered.

314. Ilium, ridge on medial surface adjacent to preacetabular notch: absent (0), present (1), strongly developed, forming a shelf (2). Ordered.

315. Ilium, preacetabulum length relative to anterior edge of pubic peduncle: reaches anteriorly to same point as (‘brachyiliac’) (0), or well past (‘dolichoiliac’) (1).

316. Ilium, depth of preacetabular process: shallow (0), deep (1).

317. Ilium, anteroventral lobe of preacetabular process: absent (0), present (1).

318. Ilium, shape of dorsal margin: convex (0), straight (1).

319. Ilium, postacetabulum length relative to ischial peduncle length: ≤ (0), > (1).

320. Ilium, depth of postacetabular process: shallow (0), deep (1).

321. Shape of posterior margin of postacetabular process of ilium: rounded, dorsal margin curves gradually into posterior margin (0), only slightly convex or straight, dorsal margin offset from posterior margin by marked posterodorsal angle (1), concave (2), tapering (3).

322. Posterior process on the dorsal end of the posterior margin of the postacetabular blade, formed by a posterior expansion of the medial brevis shelf: absent (0), present (1).

323. Obturator foramen in pubis: completely enclosed in bone (0), ventrally open notch (1), absent (2). ordered.

324. Pubic fenestra below obturator foramen: absent (0), present (1)

325. Obturator plate of ischium: largely continuous with pubic articulation (0), with large notch below the pubic peduncle, obturator process offset from pubic articulation (1) (modified from Rauhut and Pol 2019).

326. Foramen in obturator plate of ischium: absent (0), present (1) (modified from Rauhut and Pol 2019).

327. Pubis, shaft orientation: anterodorsally curved (0), straight (1), posteroventrally curved (2) (modified from Rauhut and Pol 2019).

328. Pubis, articulation between apices in adults: unfused (0), fused (1).

329. Pubis, contact between distal portions: separate distally (0), contacting (1), contacting with slit-like opening proximal to distal expansion (interpubic fenestra) (2).

330. Pubis, angle between long axes of shaft and boot: 75-90° (0), < 60° (1).

331. Pubis, morphology of symphysis: marginal (0), broad (1).

332. Pubis, morphology of obturator foramen: small and subcircular (0), large and oval (1).

333. Pubis, anterior expansion of distal end: absent (0), present (1).

334. Pubis, boot length relative to shaft length: < (0), > 30% (1), > 60% (2). Ordered.

335. Pubis, shape of boot in ventral view: broadly triangular (0), narrow, with subparallel margins (1).

336. Pubis, articulation with ilium: planoconcave (0), peg-and-socket (1).

337. Ischium, length relative to pubis length: 75-80% (0), ≤ 70% (1), > 80% (2).

338. Ischium, shaft orientation: straight (0), ventrally curved (1).

339. Ischium, articulation with ilium: planoconcave (0), peg-and-socket (1).

340. Ischium, morphology of antitrochanter: large and notched (0), reduced (1).

341. Ischium, notch ventral to obturator process: absent (0), present (1).

342. Ischium, morphology of symphysis: unexpanded (0), expanded as apron (1).

343. Ischium, cross-sectional shape of paired midshafts: oval (0), heart-shaped, medial portions of shafts extend posteriorly as midline flange (1).

344. Ischium, morphology of distal end: rounded (0), expanded, triangular (1).

345. Ischium, articulation at distal end in adults: separate (0), fused (1).

346. Femur, head orientation: 45° anteromedial (0), 10-30° anteromedial (1), medial (2). Ordered.

347. Femur, head angle: ventromedial (0), horizontal (medial) (1), dorsomedial (2). Ordered.

348. Femur, groove on proximal surface of head oriented oblique to long axis of head (‘articular groove’): absent (0), present (1).

349. Femur, oblique ligament groove on posterior surface of head: shallow, groove bounding lip does not extend past posterior surface of head (0), deep, bound medially by well-developed posterior lip (1).

350. Femur, placement of lesser trochanter relative to femoral head: does not reach ventral margin (0), rises past ventral margin (1), rises to proximal surface (2). Ordered.

351. Femur, morphology of anterolateral muscle attachments at proximal end: continuous trochanteric shelf (0), distinct lesser trochanter and attachment bulge (1).

352. Femur, development of fourth trochanter: prominent semioval flange (0), very weak or absent (1).

353. Femur, distinctly projecting accessory trochanter (derived from lesser trochanter): weak, forms slightly thickened margin of lesser trochanter (0), present as triangular flange (1).

354. Femur, M. femorotibialis externus origin medially on anterodistal surface: faint, small rugose patch (0), pronounced rugose depression that extends to distal femur (1).

355. Femur, development of medial epicondyle: rounded (0), ridge (1).

356. Femur, distal extensor groove: absent (0), present (1).

357. Femur, morphology and orientation of tibiofibularis crest: broad (0), narrow, longitudinal (1), lobular, oblique (2).

358. Femur, infrapopliteal ridge connecting medial distal condyle and crista tibiofibularis: absent (0), present (1).

359. Femur, orientation of long axis of medial condyle in distal view: anteroposterior (0), posterolateral (1).

360. Femur, projection of lateral and medial distal condyles: approximately equal (0), lateral projects distinctly further than medial, distal surface of medial is gently flattened (1).

361. Femur, morphology of distal end: central depression connected to crista tibiofibularis by a narrow groove (0), anteroposteriorly oriented shallow trough separating medial and lateral convexities (1).

362. Tibia, lateral malleolus: backs astragalus (0), overlaps calcaneum (1).

363. Tibia, shape of edge of lateral malleolus: smoothly curved (0), tabular notch (1).

364. Tibia, morphology of distal cnemial process: rounded (0), expanded proximodistally (1).

365. Tibia, morphology of lateral (fibular) condyle: large (0), small and lobular (1).

366. Tibia, anterolateral process of lateral condyle: absent or horizontal projection (0), prominent, curves ventrally (1).

367. Tibia, anteromedial buttress for astragalus: absent (0), ventral (1), marked oblique step-like ridge (2), reduced oblique ridge (3), bluntly rounded vertical ridge on medial side (4).

368. Tibia, morphology of fibular crest: narrow (0), bulbous (1).

369. Tibia, development of fibular crest: extends to proximal end of tibia as high crest (0), extends to proximal end of tibia as low ridge (1), does not extend to proximal end of tibia (2). Ordered.

370. Groove or depression on the medial side of the proximal end of the fibula: absent or only shallow concavity present (0), deep groove on the posterior half of the medial side of the proximal end, offset from anterior margin and opening at least partially posteromedially (1), large, deep depression that opens medially and is offset from the anterior margin only by a thin ridge or lip (2)

371. Ridge on the medial side of the proximal end of the fibula that extends anterodistally from the posteroproximal part: absent (0), present (1).

372. Fibula, size of iliofibularis tubercle: faint scar (0), large (1), anterolaterally curving flange (2).

373. Fibula, size of proximal end relative to width of proximal tibia: < 75% (0), ≥ 75% (1).

374. Astragalus, articulation between ascending process and fibula in adults: separate (0), fused (1).

375. Astragalus, orientation of distal condyles: ventral (0), 30-45° anterior (1).

376. Astragalus, ascending process morphology: blocky (0), laminar (1).

377. Astragalus, ascending process height relative to depth of astragalar body: less (0), subequal (1), > 1.6 times (2).

378. Astragalus, prominent proximolateral extension: absent (0), present (1).

379. Astragalus, round fossa at base of ascending process: absent (0), small (1), large (2). Ordered.

380. Astragalus, development of articular surface for distal end of fibula: large, dorsal (0), reduced, lateral (1).

381. Astragalus, posterolateral crest: absent (0), present (1).

382. Astragalus, posteromedial crest: absent (0), present (1).

383. Astragalus, articulation with calcaneum in adults: separate (0), fused (1).

384. Metatarsal I, length relative to length of metatarsal II: ≥ 50% (0), < 50% (1).

385. Metatarsal III, shape of proximal end: rectangular (0), shallow notch (1), deep notch (2). Ordered.

386. Metatarsal III, midshaft cross-sectional shape: rectangular (0), wedge-shaped, plantar surface pinched (1).

387. Metatarsal III, relative proportions of shaft: short and thick, length:transverse width ratio < 12.0 (0), long and gracile, ratio > 12.5 (1).

388. Metatarsal IV, proportions of distal end: broader than tall (0), taller than broad (1).

389. Metatarsal V, morphology of distal end: articular (0), non-articular (1).

390. Metatarsal V, length relative to length of metatarsal IV: > 50% (0), < 50% (1).

391. Antarctometatarsus: absent (0), present (1).

392. Pedal unguals, morphology of lateral and medial grooves: single (0), double (1).

393. Pedal unguals, digits III and IV, cross-sectional shape: triangular (0), elliptical (0).

394. Pedal unguals, digit II, mediolateral symmetry: symmetrical (0), asymmetrical (1).

395. Pedal digit phalanges, length of I-1 + I-2 relative to III-1: greater (0), less than or equal

SUPPLEMENTAL DATA 3-14

Files The following files are available for download in zipped file.

Suppl. Data 3. Irritator_full_matrix

Suppl. Data 4. Irritator_cranial_matrix

Suppl. Data 5. Irritator_full_matrix_equal_weighting_MPTs+strict_consensus

Suppl. Data 6. Irritator_cranial_matrix_equal_weighting_MPTS+strict_consensus

Suppl. Data 7. Irritator_full_matrix_implied_weighting_MPTs+strict

Suppl. Data 8. Irritator_full_matrix_reduced_consensus

Suppl. Data 9. Irritator_optimization_tree

Suppl. Data 10. Irritator_optimization_matrix

Suppl. Data 11. Irritator_optimization

Suppl. Data 12. Irritator_MrBayes_script

Suppl. Data 13. Irritator_jaw_movement_digital

Suppl. Data 14. Irritator_jaw_movement_print