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Published August 31, 2016 | Version v1
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Evolutionary relationships and systematics of Atoposauridae (Crocodylomorpha: Neosuchia): implications for the rise of Eusuchia

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Tennant, Jonathan P., Mannion, Philip D., Upchurch, Paul (2016): Evolutionary relationships and systematics of Atoposauridae (Crocodylomorpha: Neosuchia): implications for the rise of Eusuchia. Zoological Journal of the Linnean Society 177 (4): 854-936, DOI: 10.1111/zoj.12400, URL: http://doi.wiley.com/10.1111/zoj.12400

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urn:lsid:plazi.org:pub:0B6EFF98FF8FFFD6065B0752817DBF79
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http://publication.plazi.org/id/0B6EFF98FF8FFFD6065B0752817DBF79

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Taxonomic treatment: http://treatment.plazi.org/id/F75787E0FF9BFFC80251042180C7BE2E (URL)
Taxonomic treatment: http://treatment.plazi.org/id/F75787E0FFBCFFEE05D20100827BB900 (URL)
Figure: 10.5281/zenodo.5365884 (DOI)
Figure: 10.5281/zenodo.5365886 (DOI)
Figure: 10.5281/zenodo.5365888 (DOI)
Figure: 10.5281/zenodo.5365890 (DOI)
Figure: 10.5281/zenodo.5365892 (DOI)
Figure: 10.5281/zenodo.5365894 (DOI)
Figure: 10.5281/zenodo.5365896 (DOI)
Figure: 10.5281/zenodo.5365898 (DOI)
Figure: 10.5281/zenodo.5365900 (DOI)
Figure: 10.5281/zenodo.5365902 (DOI)
Figure: 10.5281/zenodo.5365904 (DOI)

References

  • 1. Dorsal cranial bones (skull roof, cranial table), external surface: smooth (0); ornamented (1) (Clark, 1994).
  • 2. Dorsal cranial bones (skull roof, cranial table), external surface: slightly grooved (0); heavily ornamented with deep pits and/or grooves (1); with shallow pits (2) (Clark, 1994) (character state 2 added here; added '/or' to state 1).
  • 3. Snout, external surface, sculpting: absent (0); present but to a lesser degree than cranial table (1); present, as prominent as on cranial table (2) (Gasparini et al., 2006) (character state 1 added here; added 'as prominent as on cranial table' to character state 2) [ordered].
  • 4. Rostrum, dorsal projection posterior to the external nares, relative to remainder of rostrum: absent, rostrum straight or low (0); rostrum upturned (1) (Andrade et al., 2011) (added 'posterior to the external nares, relative to remainder of rostrum').
  • 5. Skull, lateral expansion at orbits relative to rostrum: gradual (0); abrupt (1) (Clark, 1994) (added 'lateral' and 'relative to rostrum').
  • 6. Snout, lateral contour, in dorsal view: straight (0); sinusoidal ('festooned') (1) (Ortega et al., 2000).
  • 7. Snout, overall proportions: narrow oreinirostral (tall and domed) (0); nearly tubular (1); platy- rostral (broad and flat) (2) (Clark, 1994; sensu Rayfield & Milner, 2008) [ordered].
  • 8. Snout, profile of dorsal edge in lateral view (anterior to cranial table): concave (0); convex (1); approximately straight (2) (Sweetman et al., 2015).
  • 9. External nares, orientation: facing anterolaterally (0); facing dorsally (1); facing anterodorsally (2) (Clark, 1994) (character state 2 added here).
  • 10. External nares: completely divided by a septum (0); partially divided posteriorly (1); confluent, no indication of a septum (2) (Clark, 1994) [ordered].
  • 11. Orbit, orientation: facing anterolaterally (0); facing fully laterally (1) (Wilkinson, Young & Benton, 2008).
  • 13. External antorbital fenestra: large, >0.5 times the size of the orbit (0) small, ≤0.5 times the size of the orbit (1); absent (2) (Andrade et al., 2011) (changed to a multistate, adding character states 0 and 1 to replace character state 'present') [ordered].
  • 14. Antorbital fenestra, shape: rounded or dorsoventrally high (0); dorsoventrally low and anteroposteriorly elongate, slit-like (1) (Gasparini et al., 2006).
  • 15. External supratemporal fenestra: present (0); absent (1) (Ortega et al., 2006).
  • 16. External supratemporal fenestra: perforated (0); imperforated (1) (new character, adapted from Joffe, 1967).
  • 17. External supratemporal fenestra, shape: square to subrectangular (0); circular to subcircular (1); mediolaterally narrow and slit-like (2) (Andrade et al., 2011) (deleted character state 'triangular, converging medially'; added character state 2).
  • 18. External supratemporal fenestra, maximum anteroposterior length: equal to or shorter than orbits (0); longer than orbits (1) (Clark, 1994) (added 'maximum' to refine character).
  • 20. Lateral temporal fenestra in lateral view, size proportional to orbit in dorsal view: small to absent, no more than 20% of the area of the orbit (0); more than 20 to <50% of the area of the orbit (1); area is larger than 50% of the area of the orbit (2) (Andrade et al., 2011) (character state 1 added here) [ordered].
  • 21. Lateral temporal fenestra, orientation: faces laterally (0); faces dorsolaterally (1) (Andrade et al., 2011).
  • 22. Lateral temporal fenestra, shape: triangular (0); elliptical to subpolygonal (1) (Ortega et al., 2000).
  • 23. Suborbital fenestra: small, <50% of orbital area (0); between 50% and the same size as the orbit (1); larger than the orbit (2) (Andrade et al., 2011) (quantified state 1) [ordered].
  • 31. Premaxilla, maximum mediolateral width of paired premaxillae relative to that of the rostrum at the level of alveoli 4 or 5: premaxillae equal or narrower (0); rostrum narrower (1) (Jouve, 2009).
  • 32. Premaxilla, anterior to nares: narrower than, or equal to, twice the anterior nasal mediolateral width (0); broader than twice the anterior nasal width (1) (Clark, 1994) (quantified).
  • 33. Premaxilla-maxilla, distance between the anterior tip of the snout and the anterior-most position of the premaxilla-maxilla suture in dorsal view, relative to the distance between the anterior-most position of the premaxilla-maxilla suture and the posterodorsal extremity of the premaxilla in dorsal view: distance between the tip of the snout and the anterior-most position of the premaxilla-maxilla suture larger (0); distances approximately equal (1); distance between the anterior-most position of the premaxilla-maxilla suture and the posterodorsal extremity of the premaxilla larger (2) (Jouve, 2004) [ordered].
  • 34. Premaxilla-maxilla suture, small foramen in lateral surface (not for large mandibular teeth): absent (0); present (1) (Pol, 1999).
  • 35. Premaxilla, projection of the internarial bar relative to the main body of premaxilla and narial opening: does not project anterior to the main body of the premaxilla (0); strongly projected anteriorly from narial opening, extending anterior to main body of maxilla (1) (Andrade et al., 2011).
  • 36. Premaxilla, participation in internarial bar: forming at least the anterior half (0); with little participation (1) (Clark, 1994) (replaced 'ventral' with 'anterior' in character state 0).
  • 37. Premaxilla, ventral edge relative to maxilla: lower than ventral edge of maxilla, with dorsal contour of anterior part of dentary strongly concave to accommodate (0); at same height as ventral edge of maxilla (1); premaxilla ventral edge dorsal to maxilla (2) (Ortega et al., 2000) (character state 2 added here) [ordered].
  • 38. Premaxilla, perinarial crests: absent (0); present as well-defined and distinct ridges, cornering the lateral to posterior borders of the naris (1) (Andrade et al., 2011).
  • 39. Premaxilla, notch on lateral edge of external nares: absent (0); present on the dorsal half of the lateral edge of the external nares (1) (Pol, 1999).
  • 40. Premaxilla, perinarial fossa: absent (0); present (1) (Pol & Apesteguia, 2005).
  • 41. Premaxilla, postnarial fossa: absent (0); present (1) (Andrade et al., 2011).
  • 42. Premaxilla-maxilla, suture: confluent ventrally (0); opened contact on ventral edge of rostrum (1) (Clark, 1994).
  • 43. Premaxilla-maxilla contact, orientation in dorsal view, whether or not posterodorsal process is present: anteromedially directed (0); posteromedially directed (1) (Schwarz & Salisbury, 2005) (added 'whether or not posterodorsal process is present').
  • 44. Premaxilla, posterodorsal process: absent (0); present, extending posteriorly and wedging between maxillae and nasals (1) (Pol, 1999).
  • 45. Premaxilla, orientation of anterior alveolar margin: vertical (0); out-turned (1) (Sereno et al., 2001) (character state 1 modified from 'inturned').
  • 46. Maxillae, posterior palatal branches anterior to palatines: do not meet (0); meet (1) (Clark, 1994).
  • 47. Maxilla-premaxilla, suture in palatal view medial to alveolar region: sinusoidal, posteromedially directed on lateral half and anteromedially directed along medial region (0); posteromedially directed (1) (Pol, 1999) (character states 'anteromedially directed' and 'premaxillae-maxillae suture U-shaped' removed).
  • 48. Maxilla-premaxilla, lateral fossa excavating alveolus of last premaxillary tooth: absent (0); present (1) (Larsson & Sues, 2007).
  • 49. Maxilla, depression on posterolateral surface, laterally positioned: absent (0); present (1) (Wu et al., 1997) (added 'on posterolateral').
  • 52. Maxilla, ventral edge in lateral view: straight or convex (0); sinusoidal (1) (Ortega et al., 2000).
  • 53. Maxilla and premaxilla, general shape of external surface: single plane facing laterally (0); with ventral region facing laterally and dorsal region facing dorsolaterally (1) (Pol, 1999).
  • 54. Maxilla, presence of occlusal pit for reception of enlarged dentary tooth anterior to dental arcade (or M2, M stands for 'maxillary alveolus'): present (0); absent (1) (new character, adapted from Martin et al., 2014a,b).
  • 55. Maxilla, evaginated alveolar edges: absent (0); present (1) (Gasparini et al., 2006).
  • 56. Maxilla, lateral surface, unsculpted region along alveolar margin: absent (0); present (1) (Wu & Sues, 1996).
  • 57. Maxilla, sculpturing of palatal surface: absent, palatal surface smooth (0); present anteriorly, absent posteriorly (1); present throughout, palatal surface ornamented with ridges (2) (Ortega et al., 2000) [ordered].
  • 61. Maxilla, palatine process: absent (0); present, next to the anterior border of suborbital fenestrae (1) (Andrade & Bertini, 2008).
  • 64. Nasal participation in margins of external nares: present posteriorly (0); present posteriorly and medially (1) (Clark, 1994).
  • 65. Nasals: paired and unfused (0); partially or completely fused (1) (Gasparini et al., 2006) (added 'and unfused' to character state 0).
  • 66. Nasal, lateral border posterior to external nares: concave (0); straight (1); convex (2); sinusoidal (3) (character states 2 and 3 added) (Pol, 1999).
  • 67. Nasal, lateral edges: subparallel (0); oblique to one another, converging anteriorly (1) (Pol, 1999).
  • 68. Nasal, participation in antorbital fenestra: present (0); absent (1) (Ortega et al., 2000).
  • 70. Nasals, posterior mediolateral widening adjacent to the maxilla (anterior to contact with periorbital elements): abrupt (0); gradual (constant) (1); (new character, adapted from Lauprasert et al., 2011).
  • 71. Nasal-lacrimal contact: present (0); absent (1) (Clark, 1994).
  • 72. Nasal-lacrimal contact: along medial surface of lacrimal (0); lacrimal forms a point contact with nasal (1) (Clark, 1994) (changed character state 1 from 'along medial and anterior surfaces of lacrimal' to 'forms a point contact with nasal').
  • 73. Nasal, posterior tips of nasals: converge along the sagittal plane (0); separated by anterior projection of frontals (1) (Ortega et al., 2000).
  • 74. Nasal, posterior portion and anterior portion of frontal, midline anteroposterior depression: absent (0); present (1) (Montefeltro, Larsson & Langer, 2011).
  • 75. Lacrimal, total anteroposterior length relative to anteroposterior length of prefrontal: longer (0); shorter or equal to (1) (Brochu, 1999) (combined states 1 and 2).
  • 76. Lacrimal, shape: anteroposteriorly longer than mediolaterally broad (0); as anteroposteriorly long as mediolaterally broad (1) (Sereno & Larsson, 2009).
  • 77. Lacrimal, posterior extent and relationship with jugal: extending posteroventrally, widely contacting jugal (0); tapering posteroventrally, does not contact jugal or only point contact with jugal (1) (Zaher et al., 2006).
  • 78. Lacrimal and jugal, anterior margins: confluent, with no notch at the anterior contact (0); jugal edge convex, producing an anterior notch at contact (filled with maxilla) (1) (Larsson & Sues, 2007).
  • 79. Jugal, anterior extension below orbit, in dorsolateral view: does not extend beyond anterior margin of orbit (0); extends beyond anterior margin (1) (Pol, 1999).
  • 80. Jugal, dorsoventral depth of orbital portion in relation to infratemporal portion: almost the same to less than twice the depth (0); orbital portion twice the depth of the infratemporal portion (1) (Clark, 1994) (added 'to less than twice the depth' to character state 0).
  • 81. Jugal, foramen on the lateral surface near the anterior margin: absent (0); present (1) (Zaher et al., 2006).
  • 82. Jugal, anterior process length relative to infratemporal fenestrae anteroposterior length: 1.0 or less times the length (0); longer than 1.0 times the length (1) (Larsson & Sues, 2007) (changed character states 0 and 1 to define state boundary ratio).
  • 83. Jugal, orientation of base of postorbital process: directed posterodorsally (0); directed dorsally (1) (Pol, 1999).
  • 84. Jugal, location of postorbital process relative to main jugal body: anteriorly placed (0); in the middle (1); posteriorly positioned (2) (Pol, 1999) [ordered].
  • 85. Jugal portion of postorbital bar, relative to lateral surface of jugal: flush with lateral surface of jugal (0); inset (1) (Ortega et al., 2000).
  • 87. Jugal, anteroposterior ridge on lateral surface below infratemporal fenestrae: absent (0); present (1) (Pol & Norell, 2004b).
  • 88. Jugal-ectopterygoid, suture ridge: absent (0); present (1) (Montefeltro et al., 2011).
  • 89. Preorbital elements, anterior palpebral bone: no notable depression or projection (0); marked depression, developing into an incipient lateral projection (1); marked depression forming a prominent lateral projection for the support of the anterior palpebral (2) (Sereno & Larsson, 2009) (character state 0 added here) [ordered].
  • 90. Prefrontal, lateral development: reduced, no notable lateral projection (0); enlarged, extending laterally or posterolaterally over orbit (1) (Gasparini et al., 2006) (changed to 'laterally or posterolaterally' in character state 1).
  • 92. Prefrontal-frontal sutures, form paired dorsal crests: absent (0); present (1) (Pol & Powell, 2011).
  • 93. Prefrontal-lacrimal suture, crest: absent (0); present, situated anterior to orbit (1) (Andrade et al., 2011) (changed 'dorsal' to 'anterior' in character state 1).
  • 94. Prefrontal and lacrimal around orbits: forming flat rims (0); evaginated, forming elevated rims from the dorsal surface of the skull (1) (Gasparini et al., 2006).
  • 95. Prefrontal pillars (ventral process): not contacting palate (0); contacting palate (1) (Clark, 1994).
  • 96. Frontals: unfused (0); fused (1) (Clark, 1994) (changed to indicate degree of fusion).
  • 97. Frontal, mediolateral width of paired frontals between orbits: broader than nasals (0) equal or narrower than nasals (1) (Clark, 1994) (added 'equal or' to character state 1).
  • 98. Frontals, mediolateral width between orbits: narrower than posterior end (posterior end flares laterally posterior to orbits) (0); equal width or broader than posterior end (1) (new character, adapted from Sweetman et al., 2015).
  • 99. Frontal, morphology of anterior-most border of anterior process: truncated (0); wedge-like (1); bifurcated (2) (character state 2 added here) (Andrade et al., 2011) [ordered].
  • 100. Frontals, dorsal surface: flat (0); with anteroposterior ridge along midline suture (1) (Clark, 1994) (added 'along midline suture' to character state 1) [ordered] [note: see Schwarz & Salisbury (2005) for discussion of how this character relates to ontogeny].
  • 101. Frontal, dorsal anteroposterior ridge(s): restricted to the posterior portion (0); restricted to median portion (1); restricted to anterior portion (2); occupy entire length of frontal (3) (Montefeltro et al., 2011) (character state 3 added).
  • 102. Frontal, anterior extension of anterior margin: level with, or anterior to, the orbits (0); does not reach the anterior margin of the orbits (1) (Andrade et al., 2011).
  • 103. Frontal, participation in orbit border: forming great part of posterior, medial, and anterior (or anteromedial) regions (0); restricted to posterior and posteromedial region (1); restricted to medial margin (2) (Montefeltro et al., 2013) (character state 'restricted to the posterior region' removed; character states 1 and 2 added here) [ordered].
  • 104. Frontal, transverse ridge crossing anteromedial to the orbits: absent (0); present (1) (Pol et al., 2009).
  • 107. Frontal, anterior ramus with respect to the anterior tip of the prefrontal: does not extend past the anterior tip (0); extends beyond the anterior tip of the prefrontal (1) (Sereno et al., 2001) (modified states from 'ending anteriorly' and 'ending posteriorly' to remove potential gap between states).
  • 108. Frontal, lateral margin relative to the skull surface: flush (0); elevated, forming ridged orbital margins (1) (Brochu, 1999).
  • 109. Frontal, anterior process constriction with respect to main body of frontal, excluding sagittal projection into nasals anterior to orbits: absent, lateral edges parallel to subparallel (0); present, anterior portion mediolaterally constricted, with convergent lateral margins (1) (Montefeltro et al., 2013) (added 'with respect to main body of frontal, excluding sagittal projection into nasals anterior to orbits').
  • 110. Postorbital, anterolateral process: absent or poorly developed (0); well-developed, long and distally acute (1) (Clark, 1994).
  • 111. Postorbital-jugal contact, configuration: postorbital medial to jugal (0); postorbital dorsal to jugal (1) (Clark, 1994).
  • 112. Parietal-postorbital suture: absent from the dorsal surface of the skull roof (0); present on the dorsal surface of the skull roof (1) (Clark, 1994) (character broken down into characters 107-109).
  • 115. Parietal, dorsal surface: projects dorsally, relative to the skull roof (0); same level as squamosal (1); depressed relative to the squamosal (2) (Andrade et al., 2011) (character state 2 added here) [ordered].
  • 116. Parietal, posterior region dorsal surface: smooth (0); presenting a anteroposterior dorsal ridge (1); marked ventral deflection ('bevelled') in posterior portion (2); sculpted as with the rest of the skull table (3) (Montefeltro et al., 2013) (added 'anteroposterior' to character state 1; added character states 2 and 3 here).
  • 117. Parietal-squamosal emargination (anterior concavity at suture contact), posterior margin in dorsal view: absent (0); present (1) (Wilkinson et al., 2008).
  • 118. Supratemporal roof, dorsal surface: postorbital and squamosal with flat shelves extending laterally beyond quadrate contact (0); lateral edge terminating medial or immediately dorsal to medial-most point of contact with quadrate (1) (Clark, 1994) (character state 'complex' removed; character state 1 added).
  • 119. Supratemporal fenestra, medial border: flat, sculpted region (or unsculpted if rest of cranial table unsculpted) (0); forming a low sagittal crest (rims) posteriorly (1); forming a low sagittal crest (rims) along full length (2) (Clark, 1994) (character state 1 added here) [ordered].
  • 120. Supratemporal fenestra, dermal bone overhang: absent (0); present (1) (Norell, 1988).
  • 121. Supratemporal fenestra, dermal bone overhang: present only medially and posteriorly (0); present about the entire edge (1) (Norell, 1988).
  • 122. Supratemporal fenestra, shallow fossa at anteromedial corner: present (0); absent, corner smooth (1) (Brochu, 1999).
  • 126. Supratemporal fenestrae, minimum width between fenestrae, with respect to maximum width of cranial table: one-third or less of total width (0); more than one-third of total width (1) (new character; adapted from Schwarz & Salisbury, 2005).
  • 127. Orbitotemporal channel, size of the dorsal aperture: area of foramen ≤30% of that of the internal supratemporal fenestra (0); larger than 30% of the internal supratemporal fenestrae area (1) (Montefeltro et al., 2013).
  • 128. Postorbital bar between orbit and supratemporal fossa, shape: broad and solid, as broad as dorsal surface of the cranial table lateral to the supratemporal fenestra (0); much narrower (1); much narrower and connected to orbit via a thin, superficial furrow in postorbital (2) (Clark, 1994) (replaced 'with broadly sculpted dorsal surface if sculpture present' with size-related quantifier in character state 0; removed 'sculpting restricted to anterior surface' from character state 1; added character state 2) [ordered].
  • 129. Postorbital bar between orbit and supratemporal fenestra, external texture: sculpted (0); unsculpted (1) (Clark, 1994).
  • 130. Postorbital bar, lateral surface formed by: postorbital and jugal (0); only by postorbital (1) (Gasparini et al., 2006).
  • 131. Postorbital bar, shape: transversely flattened (0); transversely broad, with elliptical cross-section (1); slender and cylindrical (2) (Clark, 1994).
  • 132. Postorbital bar, shape of dorsal end nearing skull table: continuous with dorsal part of postorbital (0); dorsal part of postorbital bar constricted, distinct from the dorsal part of the postorbital (1) (Clark, 1994).
  • 133. Cranial table (skull roof), width with respect to ventral portion of skull: as wide as ventral portion of skull (quadrates covered by squamosal) (0); narrower, but still covering more than half of the mediolateral region of quadrates (1); narrower, exposing more than half of mediolateral region of quadrate (2) (Wu et al., 1997) (added 'more than half' to character states 1 and 2) [ordered].
  • 134. Squamosal and postorbital, lateral margins, dorsal view excluding the squamosal posterolateral process: parallel (0); diverging posteriorly (1); medially concave (2); converging posteriorly (3) (Ortega et al., 2000) (character state 2 added; added 'excluding the squamosal posterolateral process' to character state 0).
  • 135. Squamosal, lateral surface, longitudinal groove for attachment of the upper ear lid: absent (0); present (1) (Clark & Sues, 2002).
  • 136. Squamosal groove for upper ear lid: ventral edge is laterally displaced relative to dorsal edge (0); ventral edge is directly beneath dorsal edge (1) (Clark & Sues, 2002).
  • 137. Squamosal, dorsal edge of groove for dorsal ear lid: parallel to ventral edge (0); dorsal margin with a medial curvature (1) (Montefeltro et al., 2013).
  • 138. Squamosal, dorsolateral edge: straight and parallel to skull roof (0); bevelled ventrally, with anterolateral notch (1) (new character, adapted from Schwarz & Salisbury, 2005).
  • 139. Squamosal, posterolateral process: present (0); absent (1) (new character, adapted from Buscalioni & Sanz, 1990a).
  • 140. Squamosal, posterolateral process: level with skull table (0); depressed from skull table (1) (Sereno & Larsson, 2009).
  • 141. Squamosal, posterolateral process projection: ventrally directed, confluent with ventral rim of groove for the earflap (0); posteriorly directed and parallel to skull roof (1) (Ortega et al., 2000) (removed character state 2).
  • 142. Squamosal, posterolateral process, ornamentation: absent (0); present (1) (Larsson & Sues, 2007).
  • 143. Squamosal, posterolateral process, distal end: tapered and pointed (0); broad and rounded (1) (Larsson & Sues, 2007).
  • 144. Squamosal, anterior process extending anteriorly to the orbital margin, overlapping the postorbital, in lateral view: absent (0); present (1) (Turner & Buckley, 2008).
  • 147. Squamosal-parietal suture: flat, not elevated from the skull table (0); forms a well-developed anteroposterior groove (often bounded by elevated ridges) (1) (new character, adapted from Buffetaut, 1983).
  • 149. Quadratojugal, ornamentation at base (dorsolateral surface): absent (0); present (1) (Pol, 1999).
  • 150. Quadratojugal, length of anterior process relative to the lower temporal bar: absent or less than one-third of lower temporal bar (0); onethird to one half the length of the lower temporal bar (1); long, greater than half of the lower temporal bar (2) (Larsson & Sues, 2007) (changed character states to close gap between 'short' and 'half' of length of lower temporal bar) [ordered].
  • 151. Quadratojugal, shape of posterolateral end and relationship with quadrate: acute or rounded, tightly overlapping quadrate (0); sinusoidal ventral edge, and wide and rounded posterior edge slightly overhanging lateral surface of quadrate (1) (Pol & Norell, 2004a).
  • 153. Quadratojugal-postorbital contact, in lateral view: not in contact (0); small, point contact (1); broad contact between the quadratojugal and the posterior portion of the postorbital descending flange (2) (Clark, 1994) (added character state 0 here) [ordered].
  • 154. Infratemporal fenestra, posterior margin, dorsal view: straight (0); with an anterior projection, forming an acute angle (1) (Ortega et al., 2000) (added 'forming an acute angle' to character state 1).
  • 155. Quadrate, posterior edge: broad medial to tympanum, gently concave (0); narrow dorsal to otoccipital contact, strongly concave (1) (Clark, 1994).
  • 156. Quadrate, dorsal surface fenestration: absent (0); present (1) (Clark, 1994).
  • 157. Otic aperture (not including additional quadrate fenestrae): open posteriorly (0); closed posteriorly by quadrate and otoccipital (1) (Clark, 1994).
  • 158. Quadrate, distal body: anterior margin oriented at a right angle in relation to quadratojugal (0); anterior margin gently slopes relative to quadratojugal (1) (Montefeltro et al., 2011).
  • 159. Quadrate, pterygoid ramus: with flat ventral edge (0); with deep groove on ventral surface (1); rod-like (2) (Clark, 1994) (character state 2 added here).
  • 160. Quadrate, anterodorsal ramus in ventral view: developed, forming more than or equal to 50% of the lateral edge of the internal supratemporal fenestra (0); restricted, forming <50% of the lateral edge of the supratemporal fenestra (1) (Montefeltro et al., 2011).
  • 161. Quadrate, ventral surface: smooth, with simple muscle scars (0); with multiple developed ridges (1) (Osi € et al., 2007).
  • 162. Quadrate, condyles: with poorly developed intercondylar groove (0); medial condyle expands ventrally, being separated from the lateral condyle by a deep intercondylar groove (1) (Ortega et al., 2000).
  • 163. Quadrate, development of distal body ventral to otoccipital-quadrate contact: distinct, developing posteroventrally to contact (0); indistinct, not surpassing contact (1) (Wu et al., 1997) (added 'developing posteroventrally to contact' to character state 0, and 'not surpassing contact' to character state 1).
  • 164. Quadrate, dorsoventral height of the proximal region: ≤50% of the skull roof total width (0); more than 50% of the skull roof total width (1) (Montefeltro et al., 2013) (added 'or equal to' to character state 0).
  • 165. Cranioquadrate canal: opened laterally (0); closed laterally (1) (Clark, 1994).
  • 167. Ectopterygoid, contribution to postorbital bar: absent (0); present (1) (Sereno & Larsson, 2009).
  • 168. Ectopterygoid, main axis orientation: mediolaterally or slightly anterolaterally (0); anteroposteriorly, subparallel to anteroposterior axis of skull (1) (Pol et al., 2004) (changed laterally to mediolaterally in character state 0, and anteriorly to 'anteroposteriorly' in character state 1).
  • 169. Ectopterygoid, extent of medial projection on the ventral surface of pterygoid flanges: barely extended (0); widely extended, covering approximately the lateral half of the ventral surface of the pterygoid flanges (1) (Zaher et al., 2006).
  • 170. Ectopterygoid, anterior process: developed (0); reduced or absent (1) (Pol, 1999).
  • 171. Ectopterygoid, posterior process: developed (0); reduced or absent (1) (Pol, 1999).
  • 172. Palatines, palatal processes: do not meet on palate below narial passage (0); meet ventral to narial passage, forming part of secondary palate (1) (Clark, 1994).
  • 173. Palatine-maxilla, suture when fused at midline: palatine anteriorly rounded (0); palatine anteriorly pointed (1); suture transverse to midline axis (2) (Brochu, 1999) (character state 'palatine invaginated' removed).
  • 174. Interfenestral bar, anterior half between suborbital fenestrae, lateral margins: parallel to subparallel (0); flared anteriorly (1) (Pol et al., 2009) (added 'lateral margins').
  • 175. Interfenestral bar, posterior half between suborbital fenestrae, lateral margins: flared posteriorly (0); parallel to subparallel (1); converge posteriorly (2) (Pol et al., 2009) (added 'lateral margins'; character state 2 added here) [ordered].
  • 176. Palatines, anteroposterior axis: run parasagittally (0); diverge laterally, becoming rod-like and forming palatine bars posteriorly (1) (Martinelli, 2003).
  • 177. Palatine-pterygoid, contact on palate: run parasagittally (0); palatines firmly sutured to pterygoids (1) (Pol & Norell, 2004a).
  • 178. Pterygoids, contact with one another on palate: not in contact anterior to basisphenoid on the palate (0); pterygoids in contact (1) (Wu et al., 1997).
  • 179. Pterygoid, role of primary palate in forming choanal opening: forms posterior half of choanal opening (0); forms posterior, lateral, and part of anterior margin of choana (1); completely encloses choana (2) (Clark, 1994) [ordered].
  • 180. Pterygoid, participation in the suborbital fenestra, ventral view: forms margin of suborbital fenestra (0); excluded from suborbital fenestra by ectopterygoid-palatine contact (1) (Turner & Sertich, 2010).
  • 181. Choanae, anterior edge, location: situated between suborbital fenestrae (or anteriorly) (0); near posterior edge of suborbital fenestrae (1) (Clark, 1994).
  • 182. Choanal opening, conformation in palate: continuous with pterygoid ventral surface except for anterior and anterolateral borders (0); opens into palate through deep choanal groove (1) (Clark, 1994).
  • 183. Choanal groove: undivided (0); partially septated (1); completely septated (2) (Clark, 1994) [ordered].
  • 184. Pterygoid, quadrate process: well developed, extending posterolaterally beyond anterior margin of basioccipital (0); poorly developed, only present as an incipient projection (1) (Pol, 1999) (added 'extending posterolaterally beyond anterior margin of basioccipital' to character state 0, and 'only present as an incipient projection' to character state 1).
  • 185. Pterygoid, quadrate ramus, in ventral view: narrow and bar-like (0); broad and laminar (1) (Wu et al., 1997) (added 'bar-like' to character state 0, and 'laminar' to character state 1).
  • 186. Pterygoid, palatal surface: smooth (0); sculpted (1) (Clark, 1994).
  • 187. Pterygoid flanges: mediolaterally expanded, laterally surpassing the quadrate medial condyle (0); relatively short, and do not reach laterally to the level of the quadrate medial condyle (1) (Osi € et al., 2007).
  • 188. Basisphenoid, ventral exposure on braincase: exposed on ventral surface of braincase (0); virtually excluded from ventral surface by pterygoid and basioccipital (1) (Clark, 1994).
  • 189. Basisphenoid, lateral exposure on braincase: absent (0); present (1) (Pol, 1999).
  • 190. Basisphenoid: ventral surface continuous with surrounding bones (0); body ventrally developed and separated from the remaining elements by a posteroventral step formed by a sulcus separating it from the main occipital plane, forming a postchoanal pterygoid-basisphenoid tuberosity (1) (Montefeltro et al., 2011).
  • 191. Basisphenoid, ventral surface, mediolateral size relative to basioccipital: shorter than basioccipital (0); equal or longer than basioccipital (1) (Clark, 1994) (added 'mediolateral').
  • 192. Basioccipital: without well-developed bilateral tuberosities (0); with large, pendulous tubera (1) (Clark, 1994).
  • 193. Basioccipital, midline crest on basioccipital plate below occipital condyle: absent (0); present (1) (Turner & Sertich, 2010).
  • 194. Basioccipital and ventral part of otoccipital, orientation (when skull held horizontally): posteriorly (0); posteroventrally (1) (Gomani, 1997).
  • 195. Otoccipital, ventrolateral contact with quadrate: very narrow, otoccipital only abuts quadrate (0); broad, ventrolateral margin of otoccipital extensively contacts ventromedial portion of quadrate (1) (Clark, 1994) (added 'otoccipital only abuts quadrate' to character state 0, and 'ventrolateral margin of otoccipital extensively contacts ventromedial portion of quadrate' to character state 1).
  • 196. Supraoccipital, posterodorsal exposure in skull roof: absent (0); present (1) (Ortega et al., 2000).
  • 197. Supraoccipital, posterodorsal exposure: exposed in midline portion of posterior region of skull table (0); restricted to a thin surface attached to posterior-most portion of parietal and squamosal (1) (Montefeltro et al., 2011).
  • 198. Supraoccipital, relationship with foramen magnum: forms dorsal edge (0); otoccipitals meet dorsally, separating the foramen magnum from the supraoccipital (1) (Clark, 1994).
  • 199. Cranial nerves IX-XI, passage through braincase: all pass through common large foramen vagi in otoccipital (0); cranial nerve IX passes medial to nerves X and XI in separate passage (1) (Clark, 1994).
  • 200. Mastoid antrum, location: does not extend into supraoccipital (0); extends through transverse canal in supraoccipital to connect middle ear regions (1) (Clark, 1994).
  • 201. Mandible, outer surface sculpture, lateral surface: absent (0); present (1) (Montefeltro et al., 2011).
  • 202. Mandible, outer surface sculpture, ventral surface: present on dentary (0); present on dentary and splenial (1) (Montefeltro et al., 2011).
  • 203. Mandibular symphysis, anteroposterior length relative to mediolateral width: short, length and width subequal or shorter than wide (0); proportionally long, longer than wide (1); extremely long, length at least five times its width (2) (Andrade et al., 2011) [ordered].
  • 205. Mandibular symphysis, lateral view: shallow and tapering dorsoventrally anteriorly (0); deep and tapering dorsoventrally anteriorly (1); shallow and anterior margin convex (2) (Wu & Sues, 1996) (character state 'deep and anteriorly convex' removed).
  • 206. Mandibular symphysis, shape, in ventral view: tapering mediolaterally anteriorly, forming an angle (0); U-shaped, smoothly curving anteriorly (1); lateral edges anteroposteriorly orientated with convex anterolateral cornier and extensive, transversely orientated anterior edge (2) (Pol, 1999).
  • 207. External mandibular fenestra: absent (0); present (1) (Clark, 1994).
  • 208. External mandibular fenestra, size: present as a diminutive passage, <50% of the total size of the lateral temporal fenestra (0); present as an evident fenestra, 50% or greater than the total size of the lateral temporal fenestra (1) (Clark, 1994) (quantified both character states).
  • 209. External mandibular fenestra, orientation of main axis: horizontal to subhorizontal (0); inclined, directed anteroventrally-posterodorsally (Andrade et al., 2011) (added 'to subhorizontal' to character state 0).
  • 210. External mandibular fenestra, shape: subcircular to elliptical (0); triangular (1) (Andrade et al., 2011) (character states 'highly elliptic, anteroposterior axis much longer than dorsoventral axis, three time or more, but both ends rounded', 'slit-like, proportionally very long and both ends acute', and 'teardrop-like' removed.)
  • 211. Jaw joint, location of dorsal edge: level with or dorsolateral to occipital condyle (0); ventrolateral occipital condyle (1) (Wu & Sues, 1996).
  • 212. Dentary, lateral surface adjacent to seventh alveolus: smooth (0); with lateral concavity for reception of enlarged maxillary tooth (1) (Buckley & Brochu, 1999).
  • 213. Dentary, lateral surface below alveolar margin, at middle to posterior region of tooth row: vertically orientated, continuous with rest of lateral surface of the dentaries (0); flat surface exposed dorsolaterally, divided by ridge from the rest of the lateral surface of the dentary (1); flat, unsculpted surface confluent with rest of the lateral surface (2) (Pol & Apesteguia, 2005) (character state 2 added here).
  • 214. Dentary, relative to external mandibular fenestra: extends posteriorly beneath mandibular fenestra, posteriorly exceeding anterior margin (0); does not extend beneath fenestra, either terminating anteriorly to fenestra or only forming a point contact (1) (Clark, 1994).
  • 215. Dentary, mediolateral compression and ventrolateral surface anterior to mandibular fenestra (or of anterior portion posterior to symphysis if fenestra is absent): compressed and flat (0); uncompressed and convex (1) (Ortega et al., 1996) (added 'mediolateral').
  • 216. Dentary, sculpted below the tooth row: lacking sculpting (0); present (1) (Pol, 1999).
  • 232. Dentary, dorsolateral edge: slightly concave or straight and subparallel to anteroposterior axis of skull (0); sinusoidal, with two concave waves (1) (Ortega et al., 1996) (character state 'with single dorsal expansion and concave posteriorly' removed).
  • 233. Splenial, involvement in symphysis, in ventral view: not involved (0); involved (1) (Clark, 1994)
  • 235. Splenial, posterior to symphysis: approximately constant mediolateral thickness throughout element (0); more robust posterodorsally (1) (Ortega et al., 1996) (changed character state 'thin' to 'approximately constant mediolateral thickness throughout element'; changed character state 1 to 'more robust posterodorsally').
  • 236. Angular and posterior surangular, strong pitted pattern: absent (0); present (1); lateral surface with rugose pattern instead of pits (2) (Andrade et al., 2011) (character state 2 added here).
  • 237. Surangular, dorsal edge in lateral view: mostly straight (0); arched dorsally, excluding articular projection (1) (Clark, 1994).
  • 238. Surangular, anteroposterior ridge along the dorsolateral surface: absent (0); present (1) (Pol & Norell, 2004b).
  • 239. Surangular, extension toward posterior end of retroarticular process: along entire length (0); pinched off anterior to posterior tip (Norell, 1988).
  • 240. Articular, posterior ridge on glenoid fossa: posterior margin well developed, evidently high (0); posterior margin poorly delimited, crest absent (1) (Pol & Apesteguia, 2005).
  • 241. Articular, medial process articulating with otoccipital and basisphenoid: absent (0); present (1) (Clark, 1994).
  • 242. Retroarticular process: absent or extremely reduced (0); posteroventrally projecting and paddle-shaped (1); pointed, projects posteriorly and ventrally recurved (2); projects posteriorly and dorsally recurved (3) (Clark, 1994) (character states 'with an extensive rounded, wide, and flat (or slightly concave) surface projected posteroventrally and facing dorsomedially' and 'posteriorly elongated, triangular, and facing dorsally' removed; character states 2 and 3 added).
  • 243. Premaxillary teeth, number: five or more (0); four or fewer (1) (Wu & Sues, 1996) (character states 'six', 'three', and 'two' removed, and replaced with 'or more' and 'or fewer' in remaining character states).
  • 244. Posterior premaxillary teeth, apicobasal length: <1.5 times the size of the anterior teeth (0); 1.5 times or greater than anterior teeth (1) (Clark, 1994) (quantified and set character state boundary).
  • 245. Maxillary teeth, mesiodistal margin carinae: absent or with smooth and crenulated carinae (0); with denticulate carinae (ziphodont condition) (1) (Ortega et al., 1996) (character state 'with tubercular heterogenic denticles' removed).
  • 246. Maxillary tooth rows, middle to posterior elements: crowns not mesiodistally compressed, subcircular in cross-section (0); crowns slightly compressed mesiodistally (mesiodistal to labiolingual diameter ratio more than 0.5 at mid-height) (1); crowns highly compressed mesiodistally (mesiodistal to labiolingual diameter ratio ≤0.5 at mid-height) (2) (Pol, 1999) [ordered].
  • 247. Maxillary tooth rows, mesiodistal compression of middle to posterior elements: absent, or symmetrical compression (0); asymmetrical compression, occurring only along the distal margin giving teeth a teardrop shape (1) (Andrade & Bertini, 2008) (added 'or symmetrical compression' to character state 0).
  • 248. Maxillary teeth, middle to posterior elements, ridged ornamentation on enamel surface: absent (0); present (1) (Andrade et al., 2011).
  • 249. Maxillary teeth, enamel surface: smooth or slightly crenulated (0); with ridges at base of crown (often extending apically) (1) (Turner & Sertich, 2010).
  • 250. Maxillary teeth, striations on labial and lingual faces: present (0); absent (1) (new character, adapted from Martin et al., 2014a,b).
  • 251. Cheek teeth, base (i.e. immediately apical to root), with respect to remainder of tooth crown: not constricted (0); constricted (1) (new character, adapted from Martin et al., 2014a,b).
  • 252. Maxillary teeth, width of root with respect to crown: narrower (0); wider in anterior teeth and equal in posterior teeth (1) (Ortega et al., 2000) (changed character state 1 to 'wider in anterior teeth and equal in posterior teeth'; removed 'or equal' from character state 0).
  • 253. Maxillary teeth, posterior teeth, mediolaterally compressed lanceolate-shaped morphotype (sometimes called 'leaf-shaped'), visible in labial or lingual view, with wide crown tapering apically to a sharp point (note that the point can often be abraded): present (0); absent (1) (new character, adapted from Schwarz & Salisbury, 2005).
  • 254. Maxillary teeth, low-crowned and strongly labiolingually compressed morphotypes, forming a crown that is mesiodistally broader than it is apicobasally tall: present, apical margins orientated at <45° from horizontal (0); absent (1) (new character, adapted from Schwarz & Salisbury, 2005).
  • 255. Tooth, present at premaxilla-maxilla contact with transitional size-based morphology: absent (0); present (1) (Turner & Sertich, 2010) (added 'size-based').
  • 256. Maxillary teeth, size variation waves: absent, no tooth size variation (0); one wave of enlarged teeth (1); enlarged maxillary teeth occur in two waves (festooned) (2) (Clark, 1994).
  • 257. Enlarged maxillary teeth (at least 1.5 times the apicobasal size of remaining teeth): present at M2 and/or M3 (0); present at M4 and/or M5 (1) (Martin et al., 2014a,b). M: maxillary alveolus.
  • 261. Maxillary teeth, bulbous tooth morphotype (tribodont): present (0); absent (1) (Sweetman et al., 2015).
  • 262. Dentary teeth, anterior teeth (opposite premaxilla-maxilla contact) apicobasal length, relative to rest of dentary teeth: no more than twice the length (0); more than twice the length (1) (Clark, 1994).
  • 263. Dentary teeth, posterior teeth: occlude medial to opposing maxillary teeth (0); occlude lateral to, or interlock with, opposing maxillary teeth (1) (new character, adapted from Sweetman et al., 2015).
  • 264. Vertebrae, centra shape along axial column: cylindrical throughout (0); grade continuously from cylindrical to elongated spool-shaped (1); spool-shaped throughout (2) (Buscalioni & Sanz, 1988) (character state 1 added) [ordered].
  • 265. Cervical vertebrae: amphicoelous or amphiplatyan (0); procoelous, and posterior centrum face (condyle) with a central depression ('semi-procoely') (1); fully procoelous (2) (Clark, 1994) (character state 1 added) [ordered].
  • 267. Atlas, intercentrum size: mediolaterally wider than anteroposteriorly long (0); subequal diameters or anteroposteriorly longer (Clark, 1994).
  • 270. Cervical and dorsal vertebrae, hypapophyses or anterior keels: absent (0); present (1) (Bus- calioni & Sanz, 1988) (character states modified to present or absent).
  • 271. Dorsal vertebrae: amphicoelous or amphiplatyan (0); procoelous (1) (Clark, 1994) (replaced 'trunk' with 'dorsal').
  • 273. Posterior dorsal vertebrae, transverse process shape: dorsoventrally low and laminar (0); dorsoventrally high (1) (Buscalioni & Sanz, 1988).
  • 274. Sacral vertebrae, number: two (0); three or more (1) (Buscalioni & Sanz, 1988).
  • 275. Sacral vertebrae, orientation of transverse processes: project laterally (horizontally) (0); deflected markedly ventrally (1) (Gasparini et al., 2006).
  • 277. Caudal vertebrae: amphicoelous or amphiplatyan (0); procoelous (1) (new character, adapted from Salisbury & Frey, 2005).
  • 278. Caudal vertebrae, first: same morphology as rest of caudal series (0); biconvex (1) (new character, adapted from Salisbury & Frey, 2001).
  • 281. Coracoid, medial process: elongate posteromedial process (0); distally expanded ventromedial process (1) (Wu & Sues, 1996).
  • 282. Coracoid, distal expansion: equal to or larger than the proximal expansion (0); less expanded than the proximal region (1) (Pol et al., 2012).
  • 283. Humerus, circular depression on the posterior surface of the proximal end, for the insertion of the M. scapulohumeralis caudalis: absent (0); present (1) (Pol et al., 2012).
  • 284. Humerus, lateral and medial surfaces of distal end: flat and anteroposteriorly broad, similar in anteroposterior length to the transverse width of the distal end of the humerus (0); convex and reduced in comparison with the transverse width of the distal humerus (1) (Pol et al., 2012).
  • 287. Ulna, morphology of olecranon process: narrow and subangular (0); wide and rounded (1) (Brochu, 1999).
  • 290. Radiale, proximal end, shape: expanded symmetrically, similar to distal end (0); more expanded laterally than medially ('hatchet shaped') (1) (Buscalioni & Sanz, 1988).
  • 291. Ilium, anterior (preacetabular) process, length relative to posterior (postacetabular) process: >75% of the length of the posterior process (0); 75% or less of the length of the posterior process (1); completely absent (2) (Clark, 1994) (changed 'similar in length' to '>75% of the length of the posterior process' in character state 0; changed 'one-quarter' to '75%' in character state 1) [ordered].
  • 292. Ilium, development of the posterior (postacetabular) process: well-developed as a distinct process that extends anteroposteriorly for 60% or more of the acetabular length (0); extremely reduced or absent, extending anteroposteriorly <60% of the acetabular length (1) (Pol et al., 2012) (character state 1 changed to '<60%' to remove gap between 50 and 60%).
  • 293. Ilium, posterior end of the postacetabular process: tapering posteriorly to an acute tip (0); subrectangular with a vertically orientated posterior margin (1) (Pol et al., 2012) (removed 'with its dorsoventral height being at least 60% of the height at the origin of the postacetabular process' from character state 1).
  • 294. Pubis, shape: rod-like without expanded distal end (0); with anterodorsally-posteroventrally expanded distal end (1) (Clark, 1994) (added 'anterodorsally-posteroventrally' to character state 1).
  • 295. Pubis, anterior process: absent (0); present (1) (Clark, 1994).
  • 296. Femur, proximal development of greater trochanter: prominent, ridge-like lateral border that separates the lateral surface of the proximal femur from a flat posterior surface reaching down to the level of the fourth trochanter (0); proximodistally short trochanteric surface lacking a distinct ridge, terminating well above the fourth trochanter (1) (Pol et al., 2012).
  • 298. Tibia, distal projection of articular surfaces: medial region of distal articular surface extends further distally than the lateral region, forming a strongly oblique distal margin of the tibia (0); medial and lateral regions subequally extended, with distal margin subhorizontally orientated (1) (Pol et al., 2012).
  • 301. Astragalus, anterior margin of the tibial facet: forming a well-defined ridge that reaches medially the ball-shaped region for the articulation of metatarsals I-II and closes the proximomedial corner of the anterior hollow of the astragalus (0); forming a low ridge that is medially separated by a notch from the ball-shaped region for the articulation of the metatarsals I- II, failing to close the proximomedial corner of the anterior hollow (1) (Pol et al., 2012).
  • 305. Presacral armour: cervical and dorsal trunk shields undifferentiated, morphology grading continuously (0); cervical shields clearly differentiated from dorsal trunk shields by size and general morphology (regardless of contact between nuchal and trunk series) (1); anteriormost cervical osteoderms developed into distinct shield (2) (Andrade et al., 2011) (character state 2 added) [ordered].
  • 308. Dorsal osteoderms, shape: rounded or ovate (0); subrectangular (mediolaterally wider than anteroposteriorly long) (1); subtriangular (2); square (3) (Clark, 1994) (character state 2 added).
  • 309. Dorsal osteoderms, articular anterior process: absent (0); present (1) (Clark, 1994).
  • 310. Dorsal osteoderms, articular anterior process: as discrete convexity on anterior margin (0); well-developed process located anterolaterally ('peg and socket' articulation) (1) (Clark, 1994).
  • 315. Dorsal primary osteoderms (sensu Frey, 1988), rows: two parallel rows (0); four rows or more (1) (Clark, 1994).
  • 316. Dorsal osteoderms, accessory osteoderms (sensu Frey, 1988; i.e. osteoderms not forming part of the dorsal shield): absent (0); present (1) (Turner & Sertich, 2010).
  • 321. Dorsal osteoderms, ventral to dorsal vertebrae beneath trunk: absent (0); present (1) (Clark, 1994).
  • 322. Caudal osteoderms: absent (0) present on dorsal surface only (1); completely surrounding tail (2) (Clark, 1994) [ordered].