Published May 17, 2006
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Redescription of the Cranial Morphology of Mariliasuchus amarali, and Its Phylogenetic Affinities (Crocodyliformes, Notosuchia)
Description
Zaher, Hussam, Pol, Diego, Carvalho, Alberto B., Riccomini, Claudio, Campos, Diógenes, Nava, William (2006): Redescription of the Cranial Morphology of Mariliasuchus amarali, and Its Phylogenetic Affinities (Crocodyliformes, Notosuchia). American Museum Novitates 3512 (1): 1-40, DOI: 10.1206/0003-0082(2006)3512[1:ROTCMO]2.0.CO;2, URL: http://www.bioone.org/perlserv/?request=get-abstract&doi=10.1206%2F0003-0082(2006)3512%5B1%3AROTCMO%5D2.0.CO%3B2
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Identifiers
- LSID
- urn:lsid:plazi.org:pub:FFE0FFF89D57A810E5757544FFDE1B53
- URL
- http://publication.plazi.org/id/FFE0FFF89D57A810E5757544FFDE1B53
References
- Character definitions 1-101 were taken from Clark (1994) and have the same numeration as in the original publication. Character 5 was excluded from the analysis (because it depends on the modified definition of character 6); however, its inclusion does not affect the outcome of the analysis (except for the tree length). The additional characters are also listed here and their respective source is cited along with the character number of the original publication. Characters 1, 3, 6, 23, 37, 45, 49, 65, 67, 69, 73, 77, 79, 90, 91, 96, 97, 103, 104, 105, 107, 126, 143, 149, and 165 were set as ordered characters (marked ''+'' in this list).
- Character 1 (modified from Clark, 1994; character 1). + External surface of dorsal cranial bones: smooth (0), slightly grooved (1), and heavily ornamented with deep pits and grooves (2).
- Character 2 (modified from Clark, 1994; character 2). Skull expansion at orbits: gradual (0) or abrupt (1).
- Character 3 (modified from Clark, 1994; character 3). + Rostrum proportions: narrow oreinirostral (0) or broad oreinirostral (1) or nearly tubular (2) or platyrostral (3).
- Character 4 (Clark, 1994; character 4). Premaxilla participation in internarial bar: forming at least the ventral half (0) or with little participation (1).
- Character 5 (Clark, 1994; character 5). Premaxilla anterior to nares: narrow (0) or broad (1).
- Character 6 (modified from Clark, 1994; character 6). + External nares facing: anterolaterally or anteriorly (0), dorsally not separated by premaxillary bar from anterior edge of rostrum (1), or dorsally separated by premaxillary bar (2).
- Character 7 (Clark, 1994; character 7). Palatal parts of premaxillae: do not meet posterior to incisive foramen (0) or meet posteriorly along contact with maxillae (1).
- Character 8 (Clark, 1994; character 8). Premaxilla-maxilla contact: premaxilla loosely overlies maxilla (0) or sutured together along a butt joint (1).
- Character 9 (modified from Clark, 1994; character 9). Ventrally opened notch on ventral edge of rostrum at premaxilla-maxilla contact: absent (0) or present as a notch (1) or present as a large fenestra (2).
- Character 10 (Clark, 1994; character 10). Posterior ends of palatal branches of maxillae anterior to palatines: do not meet (0) or meet (1).
- Character 11 (Clark, 1994; character 11). Nasal-lacrimal contact: (0) or do not contact (1).
- Character 12 (Clark, 1994; character 12). Lacrimal contacts nasal along: medial edge only (0) or medial and anterior edges (1).
- Character 13 (Clark, 1994; character 13). Nasal contribution to narial border: yes (0) or no (1).
- Character 14 (Clark, 1994; character 14). Nasal-premaxilla contact: present (0) or absent (1).
- Character 15 (modified from Clark, 1994; character 15). Descending process of prefrontal: does not contact palate (0) or contacts palate (1).
- Character 16 (Clark, 1994; character 16). Postorbital-jugal contact: postorbital anterior to jugal or postorbital medial to jugal (1) or postorbital lateral to jugal (2).
- Character 17 (Clark, 1994; character 17). Anterior part of the jugal with respect to posterior part: as broad (0) or twice as broad (1).
- Character 18 (Clark, 1994; character 18). Jugal bar beneath infratemporal fenestra: flattened (0) or rod-shaped (1).
- Character 19 (Clark, 1994; character 19). Quadratojugal dorsal process: narrow, contacting only a small part of postorbital (0) or broad, extensively contacting the postorbital (1).
- Character 20 (Clark, 1994; character 20). Frontal width between orbits: narrow, as broad as nasals (0) or broad, twice as broad as nasals (1).
- Character 21 (Clark, 1994; character 21). Frontals: paired (0), unpaired (1).
- Character 22 (Clark, 1994; character 22). Dorsal surface of frontal and parietal: flat (0) or with midline ridge (1).
- Character 23 (modified from Clark, 1994; character 23; by Buckley and Brochu, 1999; character 81). + Parieto-postorbital suture: absent from dorsal surface of skull roof and supratemporal fossa (0) or absent from dorsal surface of skull roof but broadly present within supratemporal fossa (1) or present within supratemporal fossa and on dorsal surface of skull roof (2).
- Character 24 (Clark, 1994; character 24). Supratemporal roof dorsal surface: complex (0) or dorsally flat ''skull table'' developed, with postorbital and squamosal with flat shelves extending laterally beyond quadrate contact (1).
- Character 25 (modified from Clark, 1994; character 25). Postorbital bar: sculpted (if skull sculpted) (0) or unsculpted (1).
- Character 26 (modified from Clark, 1994; character 26). Postorbital bar: transversely flattened (0) or cylindrical (1).
- Character 27 (Clark, 1994; character 27). Vascular opening in dorsal surface of postorbital bar: absent (0), present (1).
- Character 28 (modified from Clark, 1994; character 28). Postorbital anterolateral process: absent or poorly developed (0) or well developed, long, and acute (1).
- Character 29 (Clark, 1994; character 29). Dorsal part of the postorbital: with anterior and lateral edges only (0) or with anterolaterally facing edge (1).
- Character 30 (Clark, 1994; character 30). Dorsal end of the postorbital bar broadens dorsally, continuous with dorsal part of postorbital (0) or dorsal part of the postorbital bar constricted, distinct from the dorsal part of the postorbital (1).
- Character 31 (Clark, 1994; character 31). Bar between orbit and supratemporal fossa broad and solid, with broadly sculpted dorsal surface (0) or bar narrow, sculpting restricted to anterior surface (1).
- Character 32 (modified from Clark, 1994; character 32). Parietal: with broad occipital portion (0) or without broad occipital portion (1).
- Character 33 (Clark, 1994; character 33). Parietal: with broad sculpted region separating fossae (0) or with sagittal crest between supratemporal fossae (1).
- Character 34 (Clark, 1994; character 34). Postparietal (dermosupraoccipital): a distinct element (0) or not distinct (fused with parietal?) (1).
- Character 35 (Clark, 1994; character 35). Posterodorsal corner of the squamosal: squared off, lacking extra ''lobe'' (0) or with unsculptured ''lobe'' (1).
- Character 36 (modified from Clark, 1994; character 36). Posterolateral process of squamosal: poorly developed and projected horizontally at the same level of the skull (0) or elongated, thin, and posteriorly directed, not ventrally deflected (1) or elongated, posterolaterally directed, and ventrally deflected (2).
- Character 37. (Clark, 1994; character 37). + Palatines: do not meet on palate below the narial passage (0) or form palatal shelves that do not meet (1) or meet ventrally to the narial passage, forming part of secondary palate (2).
- Character 38 (Clark, 1994; character 38). Pterygoid: restricted to palate and suspensorium, joints with quadrate and basisphenoid overlapping (0) or pterygoid extends dorsally to contact laterosphenoid and form ventrolateral edge of the trigeminal foramen, strongly sutured to quadrate and laterosphenoid (1).
- Character 39 (modified from Clark, 1994; character 39). Choanal opening: continuous with pterygoid ventral surface except for anterior and anterolateral borders (0) or opens into palate through a deep midline depression (choanal groove) (1).
- Character 40 (Clark, 1994; character 40). Palatal surface of pterygoids: smooth (0) or sculpted (1).
- Character 41 (Clark, 1994; character 41). Pterygoids posterior to choanae: separated (0) or fused (1).
- Character 42 (modified from Clark, 1994; character 42; by Ortega et al., 2000; character 139). Depression on primary pterygoidean palate posterior to choana: absent or moderate in size being narrower than palatine bar (0) or wider than palatine bar (1).
- Character 43 (Clark, 1994; character 43). Pterygoids: do not enclose choana (0) or enclose choana (1).
- Character 44 (modified from Clark, 1994; character 44). Anterior edge of choanae situated near posterior edge of suborbital fenestra (or anteriorly) (0) or near posterior edge of pterygoid flanges (1).
- Character 45 (Clark, 1994; character 45). + Quadrate: without fenestrae (0) or with single fenestrae (1) or with three or more fenestrae on dorsal and posteromedial surfaces (2).
- Character 46 (Clark, 1994; character 46). Posterior edge of quadrate: broad medial to tympanum, gently concave (0) or posterior edge narrow dorsal to otoccipital contact, strongly concave (1).
- Character 47 (Clark, 1994; character 47). Dorsal, primary head of quadrate articulates with: squamosal, otoccipital and prootic (0) or with prootic and laterosphenoid (1).
- Character 48 (Clark, 1994; character 48). Ventrolateral contact of otoccipital with quadrate: very narrow (0) or broad (1).
- Character 49 (Clark, 1994; character 49). + Quadrate, squamosal, and otoccipital: do not meet to enclose cranioquadrate passage (0) or enclose passage near lateral edge of skull (1) or meet broadly lateral to the passage (2).
- Character 50 (Clark, 1994; character 50). Pterygoid ramus of quadrate: with flat ventral edge (0) or with deep groove along ventral edge (1).
- Character 51 (Clark, 1994; character 51). Ventromedial part of quadrate: does not contact otoccipital (0) or contacts otoccipital to enclose carotid artery and form passage for cranial nerves IX-XI (1).
- Character 52 (Clark, 1994; character 52). Eustachian tubes: not enclosed between basioccipital and basisphenoid (0) or entirely enclosed (1).
- Character 53 (Clark, 1994; character 53). Basisphenoid rostrum (cultriform process): slender (0) or dorsoventrally expanded (1).
- Character 54 (Clark, 1994; character 54). Basipterygoid process: prominent, forming movable joint with pterygoid (0) or basipterygoid process small or absent, with basisphenoid joint suturally closed (1).
- Character 55 (modified from Clark, 1994; character 55; by Ortega et al., 2000; character 68). Basisphenoid ventral surface: shorter than the basioccipital (0) or wide and similar to or longer in length than basioccipital (1).
- Character 56 (Clark, 1994; character 56). Basisphenoid: exposed on ventral surface of braincase (0) or virtually excluded from ventral surface by pterygoid and basioccipital (1).
- Character 57 (Clark, 1994; character 57). Basioccipital: without well-developed biltaeral tuberosities (0) or with large pendulous tubera (1).
- Character 58 (Clark, 1994; character 58). Otoccipital: without laterally concave descending flange ventral to subcapsular process (0) or with flange (1).
- Character 59 (Clark, 1994; character 59). Cranial nerves IX-XI: pass through common large foramen vagi in otoccipital (0) or cranial nerve IX pass medial to nerves X and XI in separate passage (1).
- Character 60 (Clark, 1994; character 60). Otoccipital: without large ventrolateral part ventral to paroccipital process (0) or with large ventrolateral part (1).
- Character 61 (Clark, 1994; character 61). Crista interfenestralis between fenestrae pseudorotunda and ovalis nearly vertical (0) or horizontal (1).
- Character 62 (Clark, 1994; character 62). Supraoccipital: forms dorsal edge of the foramen magnum (0) or otoccipitals broadly meet dorsal to the foramen magnum, separating supraoccipital from foramen (1).
- Character 63 (Clark, 1994; character 63). Mastoid antrum: does not extend into supraoccipital (0) or extends through transverse canal in supraoccipital to connect middle ear regions (1).
- Character 64 (Clark, 1994; character 64). Posterior surface of supraoccipital: nearly flat (0), or with bilateral posterior prominences (1).
- Character 65 (modified from Clark, 1994; character 65). + One small palpebral present in orbit (0) or one large palpebral (1) or two large palpebrals (2).
- Character 66 (Clark, 1994; character 66). External nares: divided by a septum (0) or confluent (1).
- Character 67 (Clark, 1994; character 67). + Antorbital fenestra: as large as orbit (0) or about half the diameter of the orbit (1) or much smaller than the orbit (2) or absent (3).
- Character 68 (modified from Clark, 1994; character 68; by Ortega et al., 2000; character 41). Supratemporal fenestrae extension: relatively large, covering most of surface of skull roof (0) or relatively short, fenestrae surrounded by a flat and extended skull roof (1).
- Character 69 (modified from Clark, 1994; character 69). + Choanal groove: undivided (0) or partially septated (1) or completely septated (2).
- Character 70 (Clark, 1994; character 70). Dentary: extends posteriorly beneath mandibular fenestra (0) or does not extend beneath fenestra (1).
- Character 71 (modified from Clark, 1994; character 71). Retroarticular process: absent or extremely reduced (0) or very short, broad, and robust (1) or with an extensive rounded, wide, and flat (or slightly concave) surface projected posteroventrally and facing dorsomedially (2) or posteriorly elongated, triangular shaped and facing dorsally (3) or posteroventrally projecting and paddle shaped (4).
- Character 72 (Clark, 1994; character 72). Prearticular: present (0) or absent (1).
- Character 73 (modified from Clark, 1994; character 73). + Articular without medial process (0) or with short process not contacting braincase (1) or with process articulating with otoccipital and basisphenoid (2).
- Character 74 (Clark, 1994; character 74). Dorsal edge of surangular: flat (0) or arched dorsally (1).
- Character 75 (Clark, 1994; character 75). Mandibular fenestra: present (0) or absent (1).
- Character 76 (Clark, 1994; character 76). Insertion area for M. pterygoideous posterior: does not extend onto lateral surface of angular (0) or extends onto lateral surface of angular (1).
- Character 77 (modified from Clark, 1994; character 77). + Splenial involvement in symphysis in ventral view: not involved (0) or involved slightly in symphysis (1) or extensively involved (2).
- Character 78 (Clark, 1994; character 78). Posterior premaxillary teeth: similar in size to anterior teeth (0) or much longer (1).
- Character 79 (modified from Clark, 1994; character 79). + Maxillary teeth waves: absent, no tooth size variation (0) or one wave of teeth enlarged (1) or enlarged maxillary teeth curved in two waves (''festooned'') (2).
- Character 80 (Clark, 1994; character 80). Anterior dentary teeth opposite premaxilla- maxilla contact: no more than twice the length of other dentary teeth (0) or more than twice the length (1).
- Character 81 (modified from Clark, 1994; character 81). Dentary teeth posterior to tooth opposite premaxilla-maxilla contact: equal in size (0) or enlarged dentary teeth opposite to smaller teeth in maxillary tooth row (1).
- Character 82 (modified from Clark, 1994; character 82; by Ortega et al., 2000; character 120). Anterior and posterior scapular edges: symmetrical in lateral view (0) or anterior edge more strongly concave than posterior edge (1) or dorsally narrow with straight edges (2).
- Character 83 (modified from Clark, 1994; character 83; by Ortega et al., 2000; character 121). Coracoid length: up to two-thirds of the scapular length (0) or subequal in length to scapula (1).
- Character 84 (Clark, 1994; character 84). Anterior process of ilium: similar in length to posterior process (0) or one-quarter or less of the length of the posterior process (1).
- Character 85 (Clark, 1994; character 85). Pubis: rodlike without expanded distal end (0) or with expanded distal end (1).
- Character 86 (Clark, 1994; character 86). Pubis: forms anterior half of ventral edge of acetabulum (0) or pubis at least partially excluded from the acetabulum by the anterior process of the ischium (1).
- Character 87 (Clark, 1994; character 87). Distal end of femur: with large lateral facet for the fibula (0) or with very small facet (1).
- Character 88 (Clark, 1994; character 88). Fifth pedal digit: with phalanges (0) or without phalanges (1).
- Character 89 (Clark, 1994; character 89). Atlas intercentrum: broader than long (0) or as long as broad (1).
- Character 90 (modified from Clark, 1994; character 90). + Cervical neural spines: all anteroposteriorly large (0), only posterior ones rodlike (1), or all spines rodlike (2).
- Character 91 (modified from Clark, 1994; character 91; by Buscalioni and Sanz, 1988; and by Brochu, 1997; character 37 and character 7, respectively). + Hypapophyses in cervicodorsal vertebrae: absent (0) or present only in cervical vertebrae (1) or present in cervical and the first two dorsal vertebrae (2) or present up to the third dorsal vertebra (3) or up to the fourth dorsal vertebrae (4).
- Character 92 (Clark, 1994; character 92). Cervical vertebrae: amphicoelous or amphyplatian (0) or procoelous (1).
- Character 93 (Clark, 1994; character 93). Trunk vertebrae: amphicoelous or amphyplatian (0) or procoelous (1).
- Character 94 (Clark, 1994; character 94). All caudal vertebrae: amphicoelous or amphyplatian (0) or first caudal biconvex with other procoelous (1) or procoelous (2).
- Character 95 (Clark, 1994; character 95). Dorsal osteoderms: rounded or ovate (0) or rectangular, broader than long (1) or square (2).
- Character 96 (modified from Clark, 1994; character 96; and Brochu, 1997; character 40). + Dorsal osteoderms without articular anteri- or process (0) or with a discrete convexity on anterior margin (1) or with a well-developed process located anterolaterally in dorsal parasagittal osteoderms (2).
- Character 97 (modified from Clark, 1994; character 97; by Ortega et al., 2000; characters 107 and 108). + Rows of dorsal osteoderms: two parallel rows (0) or more than two (1) or more than four with ''accessory ranges of osteoderms'' (sensu Frey, 1988) (2).
- Character 98 (Clark, 1994; character 98). Osteoderms: Some or all imbricated (0) or sutured to one another (1).
- Character 99 (Clark, 1994; character 99). Tail osteoderms: dorsal only (0) or completely surrounded by osteoderms (1).
- Character 100 (Clark, 1994; character 100). Trunk osteoderms: absent from ventral part of the trunk (0) or present (1).
- Character 101 (Clark, 1994; character 101). Osteoderms: with longitudinal keels on dorsal surfaces (0) or without longitudinal keels (1).
- Character 102 (Wu and Sues, 1996; character 14). Jugal: participating in margin of antorbital fossa (0) or separated from it (1).
- Character 103 (modified from Wu and Sues, 1996; character 23). + Articular facet for quadrate condyle: equal in length to the quadrate condyles (0) or slightly longer (1) or close to three times the length of the quadrate condyles (2).
- Character 104 (modified from Wu and Sues, 1996, and Wu et al., 1997; character 24 and character 124, respectively). + Jaw joint: placed at level with basioccipital condyle (0) or below basioccipital condyle about above level of lower tooth row (1) or below level of tooth row (2).
- Character 105 (modified from Wu and Sues, 1996, and Ortega et al., 2000; character 27 and character 133, respectively). + Premaxillary teeth: five (0), four (1), three (2), or two (3).
- Character 106 (modified from Wu and Sues, 1996; character 29). Unsculptured region along alveolar margin on lateral surface of maxilla: absent (0) or present (1).
- Character 107 (modified from Wu and Sues, 1996; character 30). + Maxilla: with eight or more teeth (0) or seven (1) or six (2) or five (3) or four (4) teeth.
- Character 108 (Wu and Sues, 1996; character 33). Coracoid: without posteromedial or ventromedial process (0) or with elongate posteromedial process (1) or distally expanded ventromedial process (2).
- Character 109 (Wu and Sues, 1996; character 40). Radiale and ulnare: short and massive (0) or elongate (1).
- Character 110 (Wu and Sues, 1996; character 41). Postacetabular process: directed posteroventrally or posteriorly (0) or directed posterodorsally and much higher in position than preacetabular process (1).
- Character 111 (modified from Gomani, 1997; character 4). Prefrontals anterior to orbits: elongated, oriented parallel to anteroposterior axis of the skull (0) or short and broad, oriented posteromedially-anterolaterally (1).
- Character 112 (modified from Gomani, 1997; character 32). Basioccipital and ventral part of otoccipital: facing posteriorly (0) or posteroventrally (1).
- Character 113 (Buscalioni and Sanz, 1988; character 35). Vertebral centra: cylindrical (0) or spool shaped (1).
- Character 114 (modified from Buscalioni and Sanz, 1988; character 39). Transverse process of posterior dorsal vertebrae dorsoventrally low and laminar (0) or dorsoventrally high (1).
- Character 115 (Buscalioni and Sanz, 1988; character 44). Number of sacral vertebrae: two (0) or more than two (1).
- Character 116 (Buscalioni and Sanz, 1988; character 49). Supra-acetabular crest: present (0) or absent (1).
- Character 117 (Buscalioni and Sanz, 1988; character 54). Proximal end of radiale expanded symmetrically, similarly to the distal end (0) or more expanded proximomedially than proximolaterally (1).
- Character 118 (Ortega et al., 1996; character 5). Lateral surface of the dentary: without a longitudinal depression (0) or with a longitudinal depression (1).
- Character 119 (Ortega et al., 1996; character 9). Ventral exposure of splenials: absent (0) or present (1).
- Character 120 (Ortega et al., 1996, 2000; characters 11 and 100, respectively). Tooth margins: with denticulate carinae (0) or without carinae or with smooth or crenulated carinae (1).
- Character 121 (modified from Pol, 1999a, and Ortega et al., 2000; character 133 and character 145, respectively). Lateral surface of anterior process of jugal: flat or convex (0) or with broad shelf below the orbit with triangular depression underneath it (1).
- Character 122 (Pol, 1999a; character 134). Jugal: does not exceed the anterior margin of orbit (0) or exceeds (1).
- Character 123 (Pol, 1999a; character 135). Notch in premaxilla on lateral edge of external nares: absent (0) or present on the dorsal half of the external nares lateral margin (1).
- Character 124 (Pol, 1999a; character 136). Dorsal border of external nares: formed mostly by the nasals (0) or by both the nasals and premaxilla (1).
- Character 125 (Pol, 1999a; character 138). Posterodorsal process of premaxilla: absent (0) or present extending posteriorly wedging between maxilla and nasals (1).
- Character 126 (Pol, 1999a, and Ortega et al., 2000; character 139 and character 9, respectively). + Premaxilla-maxilla suture in palatal view, medial to alveolar region: anteromedially directed (0) or sinusoidal, posteromedially directed on its lateral half and anteromedially directed along its medial region (1) or posteromedially directed (2).
- Character 127 (Pol, 1999a; character 140). Nasal lateral border posterior to external nares: laterally concave (0) or straight (1).
- Character 128 (Pol, 1999a; character 141). Nasal lateral edges: nearly parallel (0) or entirely oblique to each other converging anteriorly (1) or oblique to each other diverging anteriorly (2).
- Character 129 (Pol, 1999a; character 143). Palatine anteromedial margin: exceeding the anterior margin of the palatal fenestrae wedging between the maxillae (0) or not exceeding the anterior margin of palatal fenestrae (1).
- Character 130 (Pol, 1999a; character 144). Dorsoventral height of jugal antorbital region respect to infraorbital region: equal or lower (0) or antorbital region more expanded than infraorbital region of jugal (1).
- Character 131 (Pol, 1999a; character 145). Maxilla-lacrimal contact: partially included in antorbital fossa (0) or completely included (1).
- Character 132 (Pol, 1999a; character 146). Lateral eustachian tube openings: located posteriorly to the medial opening (0) or aligned anteroposteriorly and dorsoventrally (1).
- Character 133 (Pol, 1999a; character 147). Anterior process of ectopterygoid: developed (0) or reduced-absent (1).
- Character 134 (Pol, 1999a; character 148). Posterior process of ectopterygoid: developed (0) or reduced-absent (1).
- Character 135 (Pol, 1999a, and Ortega et al., 2000; character 149 and character 13, respectively). Small foramen located in the premaxillo-maxillary suture in lateral surface (not for big mandibular teeth): absent (0) or present (1).
- Character 136 (Pol, 1999a; character 150). Jugal posterior process: exceeding posteriorly the infratemporal fenestrae (0) or not (1).
- Character 137 (Pol, 1999a; character 151). Compressed crown of maxillary teeth: orient- ed parallel to the longitudinal axis of skull (0) or obliquely disposed (1).
- Character 138 (Pol, 1999a; character 152). Large and aligned neurovascular foramina on lateral maxilary surface: absent (0) or present (1).
- Character 139 (modified from Pol, 1999a; character 153). External surface of maxilla and premaxilla: with a single plane facing laterally (0) or with ventral region facing laterally and dorsal region facing dorsolaterally (1).
- Character 140 (Pol, 1999a, and Ortega et al., 2000; character 154 and character 104, respectively). Maxillary teeth: not compressed laterally (0) or compressed laterally (1).
- Character 141 (Pol, 1999a; character 155). Posteroventral corner of quadratojugal: reaching the quadrate condyles (0) or not reaching the quadrate condyles (1).
- Character 142 (Pol, 1999a; character 156). Base of postorbital process of jugal: directed posterodorsally (0) or dorsally (1).
- Character 143 (Pol, 1999a; character 157). + Postorbital process of jugal: anteriorly placed (0), in the middle (1), or posteriorly positioned (2).
- Character 144 (Pol, 1999a, and Ortega et al., 2000; character 158 and character 36, respectively). Postorbital-ectopterygoid contact: present (0), absent (1).
- Character 145 (Pol, 1999a; character 161). Quadratojugal: not ornamented (0) or ornamented in the base (1).
- Character 146 (Pol, 1999a; character 162). Prefrontal-maxillary contact in the inner anteromedial region of orbit: absent (0) or present (1).
- Character 147 (Pol, 1999a; character 163). Basisphenoid: without lateral exposure (0) or with lateral exposure on the braincase (1).
- Character 148 (Pol, 1999a; character 165). Quadrate process of pterygoids: well developed (0) or poorly developed (1).
- Character 149 (modified from Pol, 1999a, and Ortega et al., 2000; character 166 and character 44, respectively). + Quadrate major axis directed: posteroventrally (0) or ventrally (1) or anteroventrally (2).
- Character 150 (Pol, 1999a; character 167). Quadrate distal end: with only one plane facing posteriorly (0) or with two distinct faces in posterior view, a posterior one and a medial one bearing the foramen aereum (1).
- Character 151 (Pol, 1999a; character 168). Anteroposterior development of neural spine in axis: well developed, covering all the neural arch length (0) or poorly developed, located over the posterior half of the neural arch (1).
- Character 152 (Pol, 1999a; character 169). Prezygapophyses of axis: not exceeding anterior edge of neural arch (0) or exceeding the anterior margin of neural arch (1).
- Character 153 (Pol, 1999a; character 170). Postzygapophyses of axis: well developed, curved laterally (0) or poorly developed (1).
- Character 154 (modified from Pol, 1999b; character 212). Shape of dentary symphysis in ventral view: tapering anteriorly forming an angle (0) or U shaped, smoothly curving anteriorly (1) or lateral edges longitudinally oriented, convex anterolateral corner, and extensive transversally oriented anterior edge (2).
- Character 155 (Pol, 1999b; character 213). Unsculpted region in the dentary below the tooth row: absent (0) or present (1).
- Character 156 (Ortega et al., 1996, and Buckley et al., 2000; character 13). Cheek teeth: not constricted at base of crown (0) or constricted (1).
- Character 157 (Ortega et al., 2000; character 42). Outer surface of squamosal laterodorsally oriented: extensive (0) or reduced and sculpted (1) or reduced and unsculpted (2).
- Character 158 (Ortega et al., 2000; character 74). Length/height proportion of infratemporal fenestra: higher than wide or equal (0) or very antero-posteriorly elongated (1).
- Character 159 (Ortega et al., 2000; character 90). Foramen intramandibularis oralis: small or absent (0) or big and slotlike (1).
- Character 160 (Ortega et al., 2000; character 146). Ectopterygoid medial process: single (0) or forked (1).
- Character 161 (modified from Gomani, 1997, and Buckley et al., 2000; character 46 and character 113, respectively). Cusps of teeth: unique cusp (0), one main cusp with smaller cusps arranged in one row (1) or one main cusp with smaller cusps arranged in more than one row (2) or several cusps of equal size arranged in more than one row (3) or multiple small cusps along edges of occlussal surface (4).
- Character 162 (Pol and Norell, 2004a; character 164). Cross section of distal end of quadrate: mediolaterally wide and anteroposteriorly thin (0) or subquadrangular (1).
- Character 163 (Pol and Norell, 2004a; character 165). Palatine-pterygoid contact on palate: palatines overlie pterygoids (0) or palatines firmly sutured to pterygoids (1).
- Character 164 (Wu et al., 1997; character 103). Squamosal descending process: absent (0) or present (1).
- Character 165 (modified from Wu et al., 1997; character 105). + Development of distal quadrate body ventral to otoccipital-quadrate contact: distinct (0) or incipiently distinct (1) or indistinct (2).
- Character 166 (Wu et al., 1997; character 106). Pterygoid flanges: thin and laminar (0) or dorsoventrally thick, with pneumatic spaces (1).
- Character 167 (Wu et al., 1997; character 108). Postorbital participation in infratemporal fenestra: almost or entirely excluded (0) or bordering infratemporal fenestra (1).
- Character 168 (Wu et al., 1997; character 109). Palatines: form margin of suborbital fenestra (0) or excluded from margin of suborbital fenestra (1).
- Character 169 (Wu et al., 1997; character 110). Angular posterior to mandibular fenestra: widely exposed on lateral surface of mandible (0) or shifted to the ventral surface of mandible (1).
- Character 170 (Wu et al., 1997; character 112). Posteroventral edge of mandibular ramus: straight or convex (0) or markedly deflected (1).
- Character 171 (modified from Wu et al., 1997; character 119). Quadrate ramus of pterygoid in ventral view: narrow (0) or broad (1).
- Character 172 (Wu et al., 1997; character 121). Pterygoids: not in contact anterior to basisphenoid on palate (0) or pterygoids in contact (1).
- Character 173 (Wu et al., 1997; character 122). Olecranon: well developed (0) or absent (1).
- Character 174 (Wu et al., 1997; character 123). Cranial table width respect to ventral portion of skull: as wide as ventral portion (0) or narrower than ventral portion of skull (1).
- Character 175 (Wu et al., 1997; character 127). Depression on posterolateral surface of maxilla: absent (0) or present (1).
- Character 176 (Wu et al., 1997; character 128). Anterior palatal fenestra: absent (0) or present (1).
- Character 177 (Pol and Norell, 2004a; character 179). Paired ridges located medially on ventral surface of basisphenoid: absent (0) or present (1).
- Character 178 (Pol and Norell, 2004a; character 180). Posterolateral end of quadratojugal: acute or rounded, tightly overlapping the quadrate (0) or with sinusoidal ventral edge and wide and rounded posterior edge slightly overhanging the lateral surface of the quadrate (1).
- Character 179 (Pol and Norell, 2004a; character 181). Orientation of quadrate body distal to otoccipital-quadrate contact in posterior view: ventrally (0) or ventrolaterally (1).
- Character 180 (Gasparini et al., 1991; character 3). Wedgelike process of the maxilla in lateral surface of premaxilla-maxilla suture: absent (0) or present (1).
- Character 181 (Pol and Norell, 2004b; character 181). Palpebrals: separated from the lateral edge of the frontals (0) or fused to each other and the lateral margin of the frontals (1).
- Character 182 (Pol and Norell, 2004b; character 182). External surface of ascending process of jugal: exposed laterally (0) or exposed posterolaterally (1).
- Character 183 (Pol and Norell, 2004b; character 183). Longitudinal ridge on lateral surface of jugal below infratemporal fenestra: absent (0) or present (1).
- Character 184 (Pol and Norell, 2004b; character 184). Dorsal surface of posterolateral region of squamosal: without ridges (0) or with three curved ridges oriented longitudinally (1).
- Character 185 (Pol and Norell, 2004b; character 185). Ridge along dorsal section of quadrate-quadratojugal contact: absent (0) or present (1).
- Character 186 (Pol and Norell, 2004b; character 186). Sharp ridge along the ventral surface of angular: absent (0) or present (1).
- Character 187 (Pol and Norell, 2004b; character 187). Longitudinal ridge along the dorsolateral surface of surangular: absent (0) or present (1).
- Character 188 (Pol and Norell, 2004b; character 188). Dorsal surface of osteoderms ornamented with anterolaterally and anteromedially directed ridges (fleur de lys pattern of Osmolska et al., 1997): absent (0) or present (1).
- Character 189 (Pol and Norell, 2004b; character 189). Cervical region surrounded by lateral and ventral osteoderms sutured to the dorsal elements: absent (0) or present (1).
- Character 190 (Pol and Norell, 2004b; character 190). Appendicular osteoderms: absent (0) or present (1).
- Character 191 (Ortega et al., 2000; character 72). Supratemporal fenestra: present (0) absent (1).
- Character 197 (Pol and Norell, 2004b; character 192). Choanal opening: opened posteriorly and continuous with pterygoid surface (0) or closed posteriorly by an elevated wall formed by the pterygoids (1).