A primitive ornithischian dinosaur from the Late Triassic of South Africa, and the early evolution and diversification of Ornithischia




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Supplementary information to

A primitive ornithischian dinosaur from the Late Triassic of South Africa, and the early evolution and diversification of Ornithischia
Richard J. Butler1,2*, Roger M. H. Smith3 and David B. Norman2

1Department of Palaeontology, The Natural History Museum, Cromwell Road, London, SW7 5BD, UK

2Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge, CB2 3EQ, UK

3Department of Karoo Palaeontology, Iziko South African Museum, PO Box 61, Cape Town 8000, South Africa
*Author for correspondence (R.Butler@nhm.ac.uk)
1. Supplementary figures 1–3

2. Measurements of SAM-PK-K8025

3. Geological Information

4. Taxonomic status of Eocursor parvus

5. Phylogenetic analysis

6. Supplementary References




1. Supplementary figures.




Figure S1. Sedimentary log of the section at Damplaats, demonstrating the stratigraphic position of the holotype specimen of Eocursor parvus (arrowed)




Figure S2. Strict consensus of two most parsimonious trees recovered by the phylogenetic analysis. Numbers beneath nodes correspond to tree description (below).





Figure S3 (see previous page). Anatomy of the holotype of Eocursor parvus (SAM-PK-K8025). (a) Basisphenoid in left lateral view. Note the presence of a large foramen on the lateral surface of the basisphenoid – a possible autapomorphy of Eocursor. (b) Preserved manual phalanges in dorsal view. Note that manual phalanges ‘a’ and ‘b’ are preserved in articulation, and that ‘b’ (the more distal phalanx) is slightly longer than ‘a’ (see below for measurements of phalanges). (c) Manual phalanges ‘a’ and ‘b’ in lateral view. Scale bars equal 10 mm. Abbreviations: accfo, accessory fossa on lateral surface of basisphenoid; bpt pr, basipterygoid process (broken); ex pt, extensor pit on dorsal surface of distal end of phalanx ‘a’; inc, canal for the internal carotid artery; mc, metacarpal; ph-a, ph-b, ph-c, phalanges ‘a’, ‘b’ and ‘c’; ps, parasphenoid.
2. Measurements of SAM-PK-K8025
Parietal

Length, frontal suture to posterior margin = 10.6 mm; width between supratemporal fenestrae = 13.5 mm.



Scapula

Height = 74 mm (right); maximum expansion, ventral = 29 mm (left); maximum expansion, dorsal = 17 mm (estimated, right); minimum anteroposterior width of shaft = 8 mm (left, right).



Humerus

Length = 67 mm (right); maximum transverse expansion, proximal end = 21 mm (right); maximum anteroposterior expansion, proximal end = 7 mm (right); maximum transverse expansion, distal end = 10.4 mm (right); maximum anteroposterior expansion, distal end: 5.7 mm (right).



Radius

Maximum transverse expansion, proximal end: 5 mm; maximum anteroposterior expansion, proximal end: 10 mm; preserved length: 27 mm; estimated length: 42 mm.



Manus (see figure S3b for explanation of phalanx labelling)

Phalanx ‘a’, length = 10 mm; phalanx ‘b’, length = 11.4 mm; phalanx ‘c’, length = 8.6 mm.



Ilium

Anteroposterior length (estimated) = 90 mm; anteroposterior length of preacetabular process (estimated) = 35 mm; maximum depth (above pubic peduncle) = 30 mm; anteroposterior length of postacetabular process = 26 mm.



Ischium

Maximum length, preserved = 97 mm (left), 93 mm (right).



Pubis

Max length = 112 mm (left); maximum length from anterior margin of obturator foramen to anterior end of prepubic process = 20 mm; max diameter of obturator foramen = 8 mm.



Femur

Length = 109 mm (right), 109 mm (left); maximum transverse expansion, proximal end = 22.7 mm (right); maximum anteroposterior expansion, proximal end = 12 mm (right); maximum transverse width, midshaft = 9.8 mm (right), 9.6 mm (left); maximum anteroposterior width, midshaft = 12.6 mm (right), 13 mm (left); maximum transverse width, distal end = 21mm (right), 23.3 (left); maximum anteroposterior width, distal end = 20 mm (right), 22 mm (left).



Tibia

Length = 138 mm (right), 136mm (left); maximum transverse expansion, proximal end = 31.5 mm (right); maximum anteroposterior expansion, proximal end = 16 mm (right); maximum transverse width, midshaft = 10 mm (left); maximum anteroposterior width, midshaft = 9.5 mm (left); maximum transverse width, distal end: 26.4 mm (right); maximum anteroposterior width, distal end = 13mm (right).



Fibula

Length = +111 mm (left); maximum transverse expansion, proximal end = 6 mm (left); maximum anteroposterior expansion, proximal end = 17 mm (left); maximum transverse width, midshaft = 4 mm (left); maximum anteroposterior width, midshaft = 5.5 mm (left); maximum transverse width, distal end = 10 mm (right); maximum anteroposterior width, distal end = 6 mm (right).



Metatarsal II

Length = 64.7 mm (right); maximum transverse expansion, proximal end = 6.7 mm (right); maximum dorsoventral expansion, proximal end = 17 mm (right); maximum transverse width, distal end: 8.7 mm (right); 8.7 mm (left); maximum dorsoventral width, distal end = 8.1mm (right); 8.2mm (left).



Metatarsal III

Length = 75 mm (right); maximum transverse expansion, proximal end = 7 mm (right); maximum dorsoventral expansion, proximal end = 14 mm (right); maximum transverse width, distal end = 9.6 mm (right), 11.6 mm (left); maximum dorsoventral width, distal end = 10.5 mm (right); 9.7 mm (left).



Phalanges (positional identity uncertain)

Lengths = 20 mm, 18 mm, 13.7 mm.




3. Geological information.
Figure S1 shows a sedimentological section of the outcrop at the Damplaats locality indicating the stratigraphic position of SAM-PK-K8025. A semi-continuous large-nodule horizon lies in the middle of the Elliot succession and marks a distinctive change in mudrock colour (see below), which is interpreted as the onset of ephemeral stream flow conditions due to climatic aridification (Smith & Kitching 1997). This nodular horizon has been previously identified as a field marker for the extinction event that defines the Triassic/Jurassic boundary in the Karoo Basin and is used as the regional datum for correlating the measured sections (Smith & Kitching 1997). Lithostratigraphically, SAM-PK-K8025 lay 5.5m below this nodular horizon in the uppermost portion of the lower Elliot Formation of Late Triassic age. The specimen is from a level 10.5 metres above an articulated skull and skeleton of Melanosaurus readi (Yates 2005) that was originally identified as Euskelosaurus sp. (Welman 1999). The specimen also lies 16.5 metres below a reworked paleosol which marks a regional hiatal surface (Smith & Kitching 1997) and that has been dated as Early Jurassic based on the abundant remains of Tritylodon in this interval (the “Tritylodon Acme Zone”; Kitching & Raath 1984).

The co-incidence of a faunal changeover occurring at a rapid sedimentological transition from warm humid to hot dry floodplain conditions suggests an environmentally induced extinction event that is interpreted as the onland equivalent of the Triassic–Jurassic faunal crisis (Bordy et al. 2004). The large plateosaurid prosauropods and the capitosaurid amphibians in the lower Elliot are widely accepted as indicator taxa of the Late Triassic (Olsen & Galton 1984) whereas the advanced cynodonts Tritylodon, Pachygenelus, and Erythrotherium are taken to indicate an Early Jurassic age for the upper Elliot (Kitching & Raath 1984). Thus it is with some certainty that the Triassic–Jurassic boundary lies in the succession between these two faunas and the distinctive change in mudrock colour will remain the most likely stratigraphic position until a method of independently dating these strata can be found.



SAM-PK-K8025 was preserved in the uppermost part of a 3 m-thick dark reddish brown massive siltstone just beneath the basal scour surface of a 1 m-thick fine-grained sandstone. Distinctive claystone-veneered lateral accretion surfaces within the sandstone display desiccation cracks and mudrock rip-up clasts indicative of flash-flood flow conditions in a small meandering river (Visser & Botha 1980). The new ornithischian specimen comprises a bundle of disarticulated but still associated post-crania mixed with disassociated skull elements. This taphonomic style is attributed to the animal having died on an open semi-arid floodplain. The carcass decomposed and fell apart without significant disturbance from scavengers. Desiccated skin possibly held the ribs and vertebrae together long enough for the skeleton to be buried, probably by windblown silt (Smith & Kitching 1997).


4. Taxonomic status of Eocursor parvus.
Eocursor parvus can be distinguished from all other basal ornithischians on the basis of several autapomorphies:
1) Accessory fossa present on lateral surface of basisphenoid, posterior to canal for internal carotid artery. A prominent accessory fossa is positioned is present on the lateral surface of the basisphenoid of Eocursor (figure S3a): this fossa is elliptical with the long-axis of the ellipse orientated anteroposteriorly and is positioned posterior to the canal for the internal carotid artery. An accessory fossa on the lateral surface of the basisphenoid is not present in other ornithischians (e.g. Galton 1989; Sereno 1991), although accessory pneumatic foramina are known in this area in theropods (Rauhut 2003).
2) Maximum transverse expansion of the distal end of the humerus is only 50% of maximum transverse expansion of the proximal end of the humerus. Compared to Eocursor parvus, the distal end of the humerus of other basal ornithischians is considerably more expanded transversely relative to the proximal end (figure 2c); the corresponding ratio is nearly 80% in Lesothosaurus (BMNH RUB17), 70-96% in Hexinlusaurus multidens (He & Cai 1984), 102% in Agilisaurus louderbacki (Peng 1992), 83% in Heterodontosaurus (Santa Luca 1980), 88% in Scutellosaurus (Colbert 1981) and 88-94% in Hypsilophodon foxii (Galton 1974).
3) Pubic obturator foramen subcircular and enlarged (maximum dorsoventral diameter of foramen is twice maximum diameter of proximal pubic shaft). In most basal ornithischians the obturator foramen is small, dorsoventrally narrow, and elliptical, with the long axis of the ellipse orientated approximately anteroposteriorly (e.g. Lesothosaurus [Sereno 1991), Heterodontosaurus [Santa Luca 1980], and Hexinlusaurus [He & Cai 1984]). By contrast, the obturator foramen is notably enlarged and subcircular in Eocursor (figure 2g).
In addition to the autapomorphies discussed above, a unique combination of characters also distinguishes Eocursor from other basal ornithischians. Comparison here is made with a range of Late Triassic and Early Jurassic ornithischians:
Pisanosaurus mertii, from the Late Triassic (Carnian) of Argentina (Bonaparte 1976), is distinguished from Eocursor by the following features: dentary teeth possess well-developed wear facets distributed systematically along the tooth row; proximal tibia lacks an accessory condyle on the anterolateral surface of the fibular condyle; only a weak notch separates the inner and fibular condyles; distal end of tibia is not strongly transversely expanded relative to the tibial shaft.
Lesothosaurus diagnosticus, from the upper Elliot Formation (Early Jurassic: Hettangian–Sinemurian) of southern Africa (Sereno 1991; Butler 2005a), can be distinguished from Eocursor on the basis of the possession of a large number of characters, including: presence of a pointed posterior parietal margin; dorsal margin of surangular medially inflected with prominent anteroposteriorly extending groove on its dorsal surface; strong expansion of the dorsal margin of the scapula; maximum manual phalangeal length less than 10% of the length of the humerus; manual phalanges lack dorsal and ventral intercondylar processes, dorsal extensor pits, and prominent collateral ligament pits; distal manual phalanges are shorter than proximal phalanges; iliac contribution to acetabulum partially closed medially by well-developed ventral flange; brevis shelf of ilium is nearly vertical; anteroposterior length of base of ischial peduncle of ilium exceeds transverse width of base of ischial peduncle; enlarged facet for the pubic peduncle of the ilium present on the proximal pubis; transverse expansion of the distal end of the femur exceeds anteroposterior expansion of the distal end of the femur.
Scutellosaurus lawleri, from the Kayenta Formation (Early Jurassic: Sinemurian–Pliensbachian) of Arizona (Colbert 1981), can be distinguished from Eocursor by possession of the following characters: anterior end of the dentary tooth row is downturned in lateral view; ilium exceeds the femur in length, dorsal margin of ilium is mediolaterally expanded into a narrow shelf; iliac contribution to acetabulum partially closed medially by well-developed ventral flange; presence of postcranial osteoderms.
Stormbergia dangershoeki, from the upper Elliot Formation (Early Jurassic: Hettangian–Sinemurian) of southern Africa (Butler 2005a), is distinguished from Eocursor by the following features: strong expansion of the dorsal margin of the scapula present; iliac contribution to acetabulum partially closed medially by well-developed ventral flange; brevis shelf of ilium is nearly vertical; anteroposterior length of base of ischial peduncle of ilium exceeds transverse width of base of ischial peduncle; shaft of ischium undergoes minimum torsion along its length; well-defined tab-like obturator process present on the ischium; ischial symphysis present at distal end of ischial shaft only.
Members of the clade Heterodontosauridae (including the unnamed heterodontosaurid from the Laguna Colorada Formation of Argentina [Báez & Marsicano 2001; Late Triassic, Norian] and Heterodontosaurus tucki and Abrictosaurus consors from the upper Elliot Formation [Early Jurassic] of southern Africa [Santa Luca 1980; Thulborn 1974]) differ from Eocursor in a number of features, including: cheek teeth are chisel-shaped with denticles restricted to the apical third of the crown; bases of crowns of cheek teeth are only very weakly expanded or are not expanded labiolingually above the root (‘cingulum’ very weak or absent); presence of dentary caniniform tooth (in Heterodontosaurus); well-developed coronoid process present; deltopectoral crest forms an angle of nearly 90 degrees with the humeral shaft; femur with rod-like fourth trochanter; distal end of fibula is reduced and split-like.

5. Phylogenetic analysis.
Eocursor parvus was previously included as an unnamed terminal taxon in a preliminary phylogenetic analysis of basal ornithischians (Butler 2005a). That analysis recovered Eocursor as a basal neornithischian, in an unresolved polytomy with Lesothosaurus diagnosticus and more derived neornithischians. Only a limited number of characters (73) were used in the analysis (Butler 2005a), and 6 characters (character numbers 7, 8, 29, 47, 50, 52) were coded incorrectly for Eocursor.

The current analysis focuses on relationships amongst basal ornithischians and includes two outgroup taxa (Marasuchus, Herrerasaurus) and 25 ingroup taxa. All operational taxonomic units are coded at the species level with the exception of Psittacosaurus, which is coded at the generic level, and the heterodontosaurid specimen BMNH A100, the specific identity of which is controversial. The majority of terminal taxa have been examined first-hand; however Marasuchus, Herrerasaurus, Pisanosaurus, Goyocephale, Homalocephale and Yinlong are coded from relevant literature. 150 morphological characters were taken from the literature and personal observations. The data matrix was constructed using the NEXUS Data Editor (http://taxonomy.zoology.gla.ac.uk/rod/NDE/nde.html); a copy of the matrix in NEXUS format is available from the lead author on request. Analyses were carried out using PAUP* v.4.0b10 (Swofford 2002); search settings used included collapsing branches with a minimum length of zero (the ‘-amb’ option), treating all characters as unordered and equally weighted, and treating multistate taxa as polymorphisms. Analysis was conducted using a heuristic search with 1000 replicates, each starting tree being produced by random stepwise addition. Three most parsimonious trees (MPTs; length = 286 steps, CI = 0.56, RI = 0.71, RC = 0.40) were recovered, differing only in the position of the problematic taxon Pisanosaurus relative to heterodontosaurids and other ornithischians. The strict consensus of these two trees is shown in supplementary figure 2.



Eocursor parvus is positioned as the sister taxon to the clade Genasauria in all three of the most parsimonious trees. The problematic clade Heterodontosauridae is recovered as the most basal group of well-known ornithischians (see also Butler 2005b), in contrast to a number of other recent studies that positioned heterodontosaurids as basal ornithopods (e.g. Sereno 1999), the sister taxon to Marginocephalia (e.g. Xu et al. 2006), or as basal neornithischians (Butler 2005a).

A Permutation Tail-Probability (PTP) test, carried out using 10,000 randomised replicates of the data set, confirms that the data set contains significant phylogenetic structure (P=0.0001). Bootstrap (10,000 replicates) and decay analyses were carried out in PAUP*, and indicate that the support for most nodes in the consensus cladogram is relatively low, although this may partially reflect morphological conservatism amongst basal ornithischian taxa. Although the clade (Abrictosaurus + (Heterodontosaurus + BMNH A100)) does not appear in the strict consensus tree (as a result of the instability of Pisanosaurus), this clade does appear in the bootstrap consensus tree, with a bootstrap support of 61%.


Tree description.

The ‘DESCRIBETREES’ option of PAUP* was used to interpret character state transformations. All transformations are based upon the strict consensus of two most parsimonious trees. Transformation was evaluated under accelerated transformation (ACCTRAN) and delayed transformation (DELTRAN) options; unambiguous synapomorphies are those that diagnose a node under both ACCTRAN and DELTRAN optimisation.

Nodes are listed in the order they appear on the strict consensus cladogram (see supplementary figure 2). Bootstrap proportions (BS) and decay indices (DI) are listed in brackets after the name of the node. Bootstrap proportions are only given for values greater than 50; decay indices are only listed for values greater than +1. Unambiguous synapomorphies are listed following the bootstrap proportions/decay indices. For simple 0-1 state changes only the character number is given; for other state changes the type of change is specified in parentheses.
1. Ornithischia (BS = 100/ DI = +5): 12, 15, 17, 48, 51, 57, 60, 67, 68 (2 to 3), 71, 72, 81, 82, 83, 84, 90, 105, 110, 120, 121, 124, 128, 130, 132, 134, 143, 145, 147
2. Heterodontosaurus + BMNH A100 (BS = 66): 70, 79
3. Eocursor parvus + Genasauria (BS = 52): 56, 64, 71 (2 to 1), 78, 97 (1 to 0), 117, 137
4. Genasauria: 101 (1 to 0), 103 (1 to 0), 104 (1 to 0), 110 (1 to 0)
5. Thyreophora (BS = 58/ DI = +2): 25, 58, 107, 130 (1 to 0), 150
6. Neornithischia: 118
7. Agilisaurus + more derived neornithischians (BS = 54): 114, 126
8. Hexinlusaurus + more derived neornithischians (BS = 76): 11, 73, 110, 112, 117 (1 to 0), 125, 127, 130 (1 to 2), 142
9. Othnieliosaurus + Cerapoda (BS = 79): 89 (1 to 2), 90 (1 to 2), 111, 131, 133
10. Cerapoda: 113, 115 (1 to 2), 143 (1 to 0)
11. Ornithopoda: 9, 37
12. Hypsilophodon + Jeholosaurus: 34, 35
13. Marginocephalia (BS = 80/ DI = +2): 9 (1 to 0), 38, 39, 68 (1 to 3), 118 (1 to 0), 122, 136, 146
14. Pachycephalosauria (BS = 64): 18, 28, 43, 44, 45, 56 (1 to 0), 96, 98, 99, 107, 116, 129
15. Homalocephale + Goyocephale (BS = 90/ DI = +2): 27, 32, 106, 108, 109, 148, 149
16. Ceratopsia: 2, 3, 7, 21, 36 (0 to 2), 42, 46 (1 to 0), 47, 127 (1 to 0)
17. Neoceratopsia: 65, 67 (1 to 0), 71 (1 to 2)
18. Chaoyangsaurus + Liaoceratops + Archaeoceratops (BS = 69/ DI = +2): 54, 78
19. Liaoceratops + Archaeoceratops (BS = 91/ DI = +3): 1 (1 to 0), 5, 22, 62
Character list and coding

The following references were used for compiling previously used characters: Maryańska & Osmólska (1985), Norman (1984), Sereno (1984, 1986, 1987, 1999, 2000), Cooper (1985), Weishampel & Heinrich (1992), Coria & Salgado (1996), Winkler et al. (1997), Zhao et al. (1999), Xu et al. (2002), Weishampel et al. (2003), You et al. (2003), Norman et al. (2004a, b), Novas et al. (2004), Butler (2005).


1. Skull, proportions: Preorbital length is 45% or more than (measured from anterior end of premaxilla/rostral to posterior end of quadrate), or (0) less than 40% (1) of skull length.

2. Rostral: Absent (0); Present (1)

3. Premaxilla, posterolateral process, length: Does not contact lacrimal (0); Contacts the lacrimal, excludes maxilla-nasal contact (1)

4. Position of the ventral (oral) margin of the premaxilla: Level with the ventral margin of the maxilla (0); Offset ventrally relative to the ventral margin of the maxilla (1)

5. Diastema between premaxilla and maxilla: Absent (0); Present (1)

6. Diastema, arched and recessed: Absent (0); Present (1)

7. Narial fossa surrounding external nares on lateral surface of premaxilla, position of ventral margin of fossa relative to the ventral margin of the premaxilla: Closely approaches the ventral margin of the premaxilla (0); Separated by a broad flat margin from the ventral margin of the premaxilla (1)

8. External naris, position of the ventral margin: Below the ventral margin of the orbit (0); Above the ventral margin of the orbit

9. Fossa positioned on premaxilla-maxilla boundary: Absent (0); Present (1)

10. Single deep elliptical fossa present along sutural line of the nasals: Absent (0); Present (1)

11. External antorbital fenestra, anteroposterior length as proportion of maximum anteroposterior length of orbit: More than 50% (0); Less than 50% (1)

12. Internal antorbital fenestra, anteroposterior length: Large, at least 15% of the skull length (0); Strongly reduced (1)

13. Antorbital fossa, shape: Triangular (0); Oval or circular (1)

14. Secondary openings (in addition to the internal antorbital fenestra) positioned anteriorly within the antorbital fossa: Absent (0); Present (1)

15. Maxilla, buccal emargination: Absent (0); Present (1)

16. Maxillary buccal emargination: Weak (0); Well-developed (1)

17. Accessory ossification present in orbital region (palpebral/supraorbital): Absent (0); Present (1)

18. Palpebral/supraorbital: Free, projects into orbit from contact with lacrimal/prefrontal (0); Incorporated into orbital margin (1)

19. Palpebral/supraorbital, number: One (0); Two (1); Three (2)

20. Exclusion of the jugal from the margin of the external antorbital fossa by lacrimal-maxilla contact: Absent (0); Present (1)

21. Widening of the skull across the jugals, chord from frontal orbital margin to extremity of jugal is more than minimum interorbital width: Absent (0); Present, skull has a triangular shape in dorsal view (1)

22. Position of maximum widening of the skull: Beneath the jugal-postorbital bar (0);

Posteriorly, beneath the infratemporal fenestra (1)

23. Jugal (or jugal-epijugal) ridge dividing the lateral surface of the jugal into two planes: Absent (0); Present (1)

24. Jugal boss: Absent (0); Present (1)

25. Cortical remodelling of the surface of at least some cranial elements: Absent (0); Present (1)

26. Cortical remodeling takes the form of a series of low nodes: Absent (0); Present (1)

27. Jugal-postorbital bar, width broader than laterotemporal fenestra: Absent (0); Present (1)

28. Jugal-postorbital joint: Elongate scarf joint (0); Short butt joint (1)

29. Jugal-squamosal contact above infratemporal fenestra: Absent (0); Present (1)

30. Jugal posterior ramus, forked: Absent (0); Present (1)

31. Postorbital: T-shaped (0); Triangular and plate-like (1)

32. Postorbital-parietal contact:Absent, or very narrow (0); Broad (1)

33. Contact between dorsal process of quadratojugal and descending process of the squamosal: Present (0); Absent (1)

34. Paraquadratic foramen or notch on the lateral (jugal) wing of the quadrate: Absent or small (0); Large (1)

35. Paraquadratic foramen: On quadrate-quadratojugal boundary (0); Within quadratojugal (1)

36. Ventral condyles of quadrate: Subequal in size (0); Medial condyle is larger (1); Lateral condyle is larger (2)

37. Frontals, proportions: Short and broad (0); Narrow and elongate (more than twice as long as wide) (1)

38. Supratemporal fenestra: Subcircular to oval (1); Anteroposteriorly elongated (1)

39. Squamosal shelf: Absent (0); Present, obscures occiput in dorsal view (1)

40. Parietal shelf: Absent (0); Present (1)

41. Parietosquamosal shelf, posterior extent: Short (0); Extended posteriorly as a frill (1)

42. Composition of the posterior margin of the parietosquamosal shelf: Parietal contributes only a small portion to the posterior margin (0); Parietal makes up at least 50% of the posterior margin (1)

43. Postorbital-squamosal bar: Bar-shaped (0); transversely broad, flattened dorsoventrally (1)

44. Postorbital-squamosal tubercle row: Absent (0); Present (1)

45. Frontal and parietal dorsoventral thickness: Thin, are not dorsoventrally thicker than other bones of the skull table such as the nasals (0); Strongly thickened dorsoventrally relative to nasals (1)

46. Paroccipital processes: Extend laterally and are slightly expanded distally (0);

Distal end pendent and ventrally extending (1)

47. Paroccipital processes, proportions: Short and deep (height ≥ 1/2 length) (0); Elongate and narrow (1)

48. Posttemporal foramen, position: Forms a notch in the dorsal margin of the OP-EX, enclosed dorsally by the squamosal (0); Entirely enclosed with the Op-Ex complex (1)

49. Basal tubera, shape: Knob-shaped (0); Plate-shaped (1)

50. Infratemporal fenestra size: Small, much smaller than orbit (0); Large, subequal or larger than the orbit (1)

51. Predentary: Absent (0); Present (1)

52. Predentary size: Shorter in length than premaxilla (0); Approximately equal in length to, or greater than, the premaxilla (1)

53. Posterior premaxillary teeth: Posterior premaxillary teeth oppose anterior dentary teeth (0); Premaxillary teeth oppose predentary only (1)

54. Tip of predentary: Flat (0); Strongly upturned (1)

55. Predentary, ventral process: Present (0); Very reduced or absent (1)

56. Dentary symphysis: V-shaped, restricted to the rostral margin of the dentary (0); Spout shaped, expanded along the ventral border of the bone (1)

57. Dentary with a marked lateral ridge in its posteior portion that demarcates an emargination forming half of the transverse width of the bone: Absent (0); Present (1)

58. Dentary tooth row (and edentulous anterior portion) in lateral view: Straight (0);

Anterior end downturned (1)

59. Ventral flange on dentary: Absent (0); Present (1)

60. Anterodorsal margin of coronoid process formed by posterodorsal process of dentary: Absent (0); Present (1)

61. Coronoid process: Absent or weak, posterodorsally oblique, depth of mandible at coronoid is less than 140% depth of mandible beneath tooth row (0); Well-developed, distinctly elevated, depth of mandible at coronoid is more than 180% depth of mandible beneath tooth row (1)

62. Coronoid process, position: Posterior to dentition (0); Lateral to dentition (1)

63. External mandibular fenestra, situated on dentary-surangular-angular boundary: Present (0); Absent (1)

64. Anteroposteriorly extended ridge on lateral surface of surangular, anterior to jaw joint: Absent (0); Present (1)

65. Retroarticular process: Elongate (0); Short or absent (1)

66. Node-like cortical remodelling of the lateral surface of the dentary and surangular: Absent (0); Present (1)

67. Level of jaw joint relative to maxillary tooth row: Level, not offset ventrally (0); Offset ventrally (1)

68. Premaxillary teeth, number: Six (0); Five (1); Four (2); Three (3); Two (4); Premaxillary teeth absent (5)

69. Premaxillary teeth, crown expanded above root: Crown is unexpanded mesiodistally above root, no distinction between root and crown is observable (0); Crown is at least moderately expanded above root (1)

70. Premaxillary teeth increase in size posteriorly: Absent, all premaxillary teeth subequal in size (0); Present, posterior premaxillary teeth are significantly larger in size than anterior teeth (1)

71. Maxillary & dentary crowns, shape: Apicobasally tall, blade-like (0); Apicobasally short & triangular (1); Chisel-shaped, denticles restricted to apical third of crown (2)

72. Maxillary/dentary teeth, marginal ornamentations: Fine serrations set at right angles to the margin of the tooth (0); Coarse serrations (denticles) angle upwards at 45 degrees from the margin of the tooth (1)

73. Apicobasally extending ridges on lingual/labial surfaces of maxillary/dentary teeth: Absent (0); Present (1)

74. Apicobasally extending ridges, form: Some apicobasally extending ridges present, but do not correspond to the number of marginal denticles (0); Apicobasally extending denticles confluent with marginal denticles (1)

75. Prominent primary ridge on labial side of maxillary teeth: Absent (0); Present (1)

76. Prominent primary ridge on lingual side of dentary teeth: Absent (0); Present (1)

77. Position of maxillary/dentary primary ridge: Centre of the crown surface, giving the crown a relatively symmetrical shape in lingual/labial view (0); Offset, giving crown asymmetrical appearance (1)

78. At least moderately developed labiolingual expansion of crown (‘cingulum’) on maxillary/dentary teeth: Present (0); Absent (1)

79. Heterodont dentary dentition: No substantial heterodonty is present in dentary dentition (0); Single, enlarged, caniform anterior dentary tooth, crown is not mesiodistally expanded above root (1); Anterior dentary teeth are strongly recurved and caniform, but have crowns expanded mesiodistally above their roots and are not enlarged relative to other dentary teeth (2).

80. “Special” replacement foramina medial to dentary and maxillary tooth rows: Absent (0); Present (1)

81. Recurvature in maxillary and dentary teeth: Present (0); Absent (1)

82. Overlap of adjacent crowns in maxillary and dentary tooth rows: Absent (0); Present (1)

83. Crown is mesiodistally expanded above root in cheek teeth: Absent (0); Present (1)

84. Position of maximum apicobasal crown height in dentary/maxillary tooth rows: Anterior portion of tooth rows (0); Central portion of tooth rows (1)

85. Spaces between alveolar border and crowns of adjacent functional teeth in maxillary/dentary tooth rows: Absent (0); Present (1)

86. Epipophyses on anterior cervicals: Present (0); Absent (1)

87. Articulation between the zygapophyses of dorsal vertebrae: Flat (0); Tongue-and-groove (1)

88. Dorsals, number: 12-13 (0); 15 (1)

89. Sacrals, number: Two (0); Four or five (1); Six or more (2)

90. Number of sacral ribs contacting main body of ilium (posterior to preacetabular process): Two or three (0); Four (1); Five (1)

91. Sacrum, accessory articulation with anterior prepubis: Absent (0); Present (1)

92. Posterior sacral ribs are considerably longer than anterior sacral ribs: Absent (0); Present (1)

93. Ossified sternal ribs: Absent (0); Present (1)

94. Sternal plates, shape: Kidney-shaped (0); Shafted or hatchet-shaped (rod-like posterolateral process, expanded anterior end) (1)

95. Proportions of humerus and scapula: Scapula longer than, or subequal to, the humerus (0); Humerus substantially longer than the scapula

96. Deltopectoral crest development: Well-developed, projects anteriorly as a distinct flange (0); Rudimentary, is at most a thickening on the anterolateral margin of the humerus (1)

97. Angle between distal deltopectoral crest and shaft of humerus: Less than 90 degrees (0); About 90 degrees (1)

98. Humeral length: More than 60% of femoral length (0); Less than 55% of femoral length (1)

99. Humeral shaft form, in anterior or posterior view: Relatively straight (0); Strongly bowed laterally along length (1)

100. Longest manual phalanx as percentage of length of humerus: Less than 10% (0);

More than 15% (1)

101. Metacarpals with block-like proximal ends: Absent (0); Present (1)

102. Penultimate phalanx of the second and third manual digits: Shorter than first phalanx (0); Longer than the first phalanx (1)

103. Extensor pits on the dorsal surface of the distal end of at least some manual phalanges/metacarpals: Absent or poorly developed (0); Deep, well-developed (1)

104. Manual unguals strongly recurved with prominent flexor tubercle: Absent (0);

Present (1)

105. Preacetabular process of ilium, shape and length: Short, tab-shaped, distal end is posterior to pubic peduncle (0); Elongate, strap-shaped, distal end is anterior to pubic peduncle (1)

106. Preacetabular process of ilium, lateral deflection in dorsal view: 10-20 degrees (0); More than 30 degrees (0)

107. Dorsal margin of preacetabular process and dorsal margin of ilium above acetabulum: Narrow, not transversely expanded (0); Dorsal margin is transversely expanded to form a narrow shelf (1)

108. Preacetabular process of the ilium distally expanding in dorsal view: Absent (0); Present (1)

109. Subtriangular process extending medially from the dorsal margin of the iliac blade: Absent (0); Present (1)

110. Brevis shelf & fossa: fossa faces ventrolaterally and visible in lateral view along entire length, creating a deep postacetabular portion (0); fossa faces ventrally and posterior portion cannot be seen in lateral view, forming at least a narrow horizontal shelf (1)

111. Medioventral flange of ilium, partially closes the acetabulum: Present (0); Absent (1)

112. Supra-acetabular ‘flange’: Present (0); Absent (1)

113. Ischial peduncle of the ilium: Projects ventrally (0); Broadly swollen, projects ventrolaterally (1)

114. Pubic peduncle of ilium: Large, elongate, robust (0); Reduced in size, shortened relative to ischial peduncle (1)

115. Length of the postacetabular process as a % of the ilium length: 20% or less (0); Around 30% (1); 40% or more (2)

116. Ischium, shape of shaft in lateral view: Relatively straight (0); Gently curved along length

117. Dorsal groove on the ischium: Absent (0); Present (1)

118. Tab-shaped obturator process on ischium: Absent (0); Present (1)

119. Ischial symphysis, length: Ischium forms a median symphysis with the opposing blade along at least 50% of its length (0); Ischial symphysis present distally only (1)

120. Pubis, orientation: Anteroventral (0); Rotated posteroventrally to lie alongside the ischium (opisthopubic) (1)

121. Shaft of pubis (postpubis), shape: Blade-shaped (0); Rod-shaped (1)

122. Shaft of pubis (postpubis), length: Approximately equal in length to the ischium (0); reduced in length (1)

123. Reduction of pubic shaft: extends for approximately half the length of the ischium (0); Very short or absent (1)

124. Prepubic process: Absent (0); Present (1)

125. Prepubic process: Compressed mediolaterally, dorsoventral height exceeds mediolateral width (0); Rod-like, mediolateral width exceeds dorsoventral height (1)

126. Prepubic process, length: Stub-like and poorly developed, extends only a short distance anterior to the pubic peduncle of the ilium (0); Elongated into distinct anterior process (1)

127. Prepubic process: does not extend beyond distal end of preacetabular process of ilium (0); Extends beyond distal end of preacetabular process of ilium (1)

128. Extent of pubic symphysis: Elongate (0); Restricted to distal end of pubic blade, or absent (1)

129. Femur/tibia ratio: Femur shorter than tibia (0); Tibia shorter than femur (1)

130. Greater or ‘dorsolateral’ (Langer, 2003) trochanter: 'Dorsolateral' trochanter is low proximodistally extending ridge only (0); 'Dorsolateral' trochanter expanded anteriorly as distinct flange, but is not anteroposteriorly broader than the anterior trochanter (1); 'Dorsolateral' trochanter expanded strongly anteriorly and proximally, exceeds anteroposterior width of anterior trochanter (2)

131. Femoral head: Confluent with greater trochanter, fossa trochanteris is groove-like (0); Fossa trochanteris is modified into distinct constriction separating head and greater trochanter (1)

132. Form of anterior trochanter: Trochanteric shelf either merging with the proximal femur, or ending in a small, pointed, spike (0); Broadened, prominent, ‘wing’ or ‘blade’ shaped (1)

133. Level of most proximal point of anterior trochanter relative to level of proximal femoral head: Anterior trochanter is positioned distally on the shaft, and separated from dorsolateral trochanter/greater trochanter by deep notch visible in medial view (0); Anterior trochanter positioned proximally, approaches level of proximal surface of femoral head, closely appressed to dorsolateral/greater trochanter (no notch visible in medial view) (1)

134. Fourth trochanter, pendent: Absent (0); Present (1)

135. Fourth trochanter, position: Located entirely on proximal half of femur (0); Positioned at midlength, or distal to midlength (1)

136. Anterior (extensor) intercondylar groove: Absent (0); Present (1)

137. Maximum expansion of distal tibia relative to proximal: Distal tibia is considerably less expanded than proximal (0); Maximum expansion of distal tibia is subequal to that of proximal tibia (1).

138. Distal tibia: Subquadrate, posterolateral process is not substantially developed (0); Elongate caudolateral process, backs fibula (1)

139. Fibula reduced to narrow splint distally: Absent (0); Present (1)

140. Fibular facet on the proximolateral surface of the astragalus: Large (0); Reduced to small articulation (1)

141. Calcaneum: Facet for tibia absent (0); Well-developed facet for tibia present (1)

142. Medial distal tarsal : Articulates distally with metatarsal 3 only (0); Articulates distally with metatarsals 2 and 3 (1).

143. Digit 1: Metatarsal 1 robust and well-developed, distal end of phalanx 1-1 projects beyond the distal end of metatarsal 2 (0); Metatarsal 1 reduced & proximally splint like, end of phalanx 1-1 does not extend beyond the end of metatarsal 2 (1)

144. Proximal end of metatarsal 2, anteroposterior expansion relative to proximal end of metatarsal 3: Anteroposterior expansion of metatarsal 2 is less than anteroposterior expansion of metatarsal 3 (0); Anteroposterior expansion of metatarsal 2 is more than anteroposterior expansion of metatarsal 3 (1).

145. Metatarsal 5, length: Length is more than 50% of the length of metatarsal 3 (0); Less than 25% of the length of metatarsal 3

146. First phalanx of digit 1 is the longest non-ungual phalanx of the pes: Absent (0); Present (1)

147. Epaxial ossified tendons: Absent (0); Present (1)

148. Ossified hypaxial tendons, present on caudal vertebrae: Absent (0); Present (1)

149. Ossified tendons, arrangement: Longitudinally arranged throughout vertebral column (0); Basket-like arrangement in caudal region (1)



150. Postcranial dermal osteoderms: Absent (0); Present (1)
Specimens and references used for coding operational taxonomic units
Institutional abbreviations: BMNH, Natural History Museum, London, UK; BRSMG, Bristol City Museum and Art Gallery, Bristol, UK; BP, Bernard Price Institute for Palaeontological Research, Johannesburg, South Africa; BYU, Earth Science Museum, Brigham Young University, Provo, Utah, USA; CAMSM, Sedgwick Museum, University of Cambridge, Cambridge, UK; GZG, Geowissenschaftliches Zentrum der Universität Göttingen, Göttingen, Germany; IGCAGS, Institute of Geology, Chinese Academy of Sciences, Beijing, People’s Republic of China; IVPP, Institute of Vertebrate Palaeontology and Palaeoanthropology, Bejing; MNA, Museum of Northern Arizona, Flagstaff, USA; MOR, Museum of the Rockies, Bozeman, Montana, USA; SAM-PK, South African Museum (Iziko Museums of Cape Town), Cape Town, South Africa; SGWG, Sektion Geologische Wissenschaften Greifswald, Ernst-Moritz Universität, Greifswald, Germany; UCMP, University of California Museum of Paleontology, Berkeley, USA; ZDM, Zigong Dinosaur Museum, Dashanpu, People’s Republic of China.
Marasuchus liloensis: Bonaparte (1975), Sereno & Arcucci (1994).

Herrerasaurus ischigualastensis: Novas (1993), Sereno (1993), Sereno & Novas (1993); MCZ 7063, 7064.

Abrictosaurus consors: Thulborn (1974), Hopson (1975); BMNH RUB54 (holotype).

Agilisaurus louderbacki: Peng (1992), Barrett et al. (2005); ZDM 6011 (holotype).

Archaeoceratops oshimai: Dong & Azuma (1997), You & Dodson (2003); IVPP V11114 (holotype); IVPP V11115 (paratype).

Chaoyangsaurus youngi: Zhao et al. (1999); IGCAGS V371 (holotype).

Emausaurus ernsti: Haubold (1990); SGWG 85 (holotype).

Goyocephale lattimorei: Perle et al. (1982).

Heterodontosaurus tucki: Santa Luca (1980); SAM-PK-K337 (holotype); SAM-PK-K1332 (referred specimen).

Hexinlusaurus multidens: He & Cai (1984), Barrett et al. (2005); ZDM T6001 (holotype).

Homalocephale calathocercos: Maryańska & Osmólska (1974).

Hypsilophodon foxii: Galton (1974); BMNH R197 (holotype); other BMNH specimens (see Galton 1974).

Jeholosaurus shangyuanensis: Xu et al. (2000); IVPP V12529 (holotype); IVPP V12530.

Lesothosaurus diagnosticus: Thulborn (1970, 1972), Santa Luca (1984), Sereno (1991), Butler (2005a); BMNH RUB17, RUB23 (syntypes); BMNH R11956, R8501 (referred specimens); SAM-PK-K400, K401, K1106 (referred specimens).

Liaoceratops yanzigouensis: Xu et al. (2002); IVPP V12738 (holotype).

Lycorhinus angustidens: Gow (1975, 1990), Hopson (1975); SAM-PK-K3606 (holotype); BP/1/4244, BP/1/5253 (referred, Gow 1990).

Othnieliosaurus celer: Galton & Jensen (1973), Galton (1983), Galton (2007); BYU ESM-163R.

Orodromeus makelai: Scheetz (1999); MOR 294 (holotype), MOR 403, 473, 623, 1141 (referred specimens).

Pisanosaurus mertii: Bonaparte (1976), Sereno (1991).

Psittacosaurus: Sereno (1987, 1990a, 1990b), Sereno & Chao (1988), Sereno et al. (1988); IVPP V738, 740-1, 749 (Psittacosaurus sinensis holotype and referred specimens); IVPP V7705 (Psittacosaurus meileyingensis holotype).

Scelidosaurus harrisonii: Owen (1861, 1863), Norman et al. (2004a); BMNH R1111 (lectotype); BMNH R5909, R6704; BRSMG Ce12785; CAMSM X 39256.

Scutellosaurus lawleri: Colbert (1981), Rosenbaum & Padian (2000); MNA P1.175 (holotype); MNA P1.1752 (paratype); UCMP 130580, 130581, 170829, 175166, 175167, 175168 (referred specimens).

Stenopelix valdensis: Sues & Galton (1982), Sereno (1987); GZG 741/2 (holotype).

Stormbergia dangershoeki: Butler (2005); SAM-PK-K1105 (holotype); BMNH R11000 (paratype); BP/1/4885 (referred).

Yinlong downsi: Xu et al. (2006).

Wannanosaurus yansiensis: Hou (1977); IVPP V4447 (holotype), IVPP V4447.1 (paratype).
Character scorings
Marasuchus lilloensis

???000???? ????0????? ?????????? ?????????? ?????????? ?????????? ?????????? 000-0?-0?? 0000010?00 00???0?00? ????00000- 0000100000 00-0---000 0000000000 010?0?00-0



Herrerasaurus ischigualastensis

0000001100 00100-0--0 0-000-0001 0000010000 --00000001 0----00000 0000000200 000-00-000 0000000100 000??01101 1111000000 0000100000 00-0---010 0000000000 00000000-0


Abrictosaurus consors

00?111000? 010?10100? 0-??0-???? ?????????? ?????????0 1010101001 1000?01300 210-0?-00? 11110????? ?0???0?00? ??1?100001 ?1001????? ????????01 ?1?100???? ??1?1????0


Agilisaurus louderbacki

0000000001 0100111010 0-000-0000 00?0010000 --00010?00 1000?11001 1010001110 210-00-121 111101?111 000?10010? ????100000 0001101101 10-1010101 010100110? 1011101000


Archaeoceratops oshimai

0110101000 0100111000 1110111010 10???20111 1?00001?00 1111011001 11?1110310 2111111001 11111?1022 10???????? ????100001 11112??011 11111101?2 111????101 1101111000


BMNH A100

?0??110??? 010?11???? 0-0?0-0??? ???????0?? ?????????? ?????0?0?? ???????301 210-00-110 11110????? ?????????? ?????????? ?????????? ?????????? ?????????? ??????????


Chaoyangsaurus youngi

11?000110? ?1??11???? 100011???0 ???002???? ????????0? 1?11011001 1010110410 211010100? 11110????? ?????????? ?????????? ?????????? ?????????? ?????????? ??????????


Emausaurus ernsti

000000000? 0100111000 0-00100001 00000?0?00 --000???00 ??0??11101 0001001110 110-00-101 11110????? ?????????? 0????????? ?????????? ?????????? ?????????? ?????????1


Eocursor parvus

?????????? ?????????? ?????????? ???????000 --??0????? ?????11001 0001001??? 110-?0-1?? 1?1?0?0?11 0???000001 ?11?100001 1000101001 10-100-101 010100110? ???1?????0


Goyocephale lattimorei

?0011100?0 ?1??11111? 0-?0111100 -1????0011 0011110?1? ?0???0?001 101?01?311 110?00-11? 11110?1?22 ?101?1011? ????111111 11??2????? ?????????? ???????1?? ?1????1110


Heterodontosaurus tucki

0011110001 0101111000 0-010-010- 0000020000 --00000101 1010101001 1000001301 211011P010 100110?021 ?00?001001 1111100001 11001000?1 10-100-102 011100011? ?01?101000


Hexinlusaurus multidens

??0??????1 1100111001 0-000-0000 00?00?0000 --00010??0 ?????11001 10?0?01??? 211000-101 111101?111 000?100000 0000100001 0101100111 10-1111102 010100110? 1111101000


Homalocephale calathocercos

?????????? 11--11111- 0-10111100 -110000011 0011110?10 ?????????? ??????1??? 11??0??1?? 11110?1?22 1101?????? ????111111 1111210011 11111111?? 111?0??101 11????1110


Hypsilophodon foxii

000110001? 111111100P 0-000-0000 0011101000 --00010100 1110011001 1010001110 2110010101 1111010122 1010000000 0000100001 1111200111 10-1111102 1111001101 1101101100


Jeholosaurus shangyuanensis

00?0000011 111?11???1 0-000-0000 0011101000 --00010?00 1110011001 1010?010?0 11110?-1?? 111101???? ?????????? ?????????? ?????????? ?????????2 111?001101 111110????


Lesothosaurus diagnosticus

000000000? 0100101000 0-000-00?? 00000?0000 --00000100 1000011001 0001001010 110-00-101 1111000?11 0???000100 0000100000 0000101001 10-100-101 0101001101 10111?1000


Liaoceratops yanzigouensis

0110101000 010011???0 11100-1010 1010020111 1100001100 1111011011 1110110310 2111111001 11110????? ?????????? ?????????? ?????????? ?????????? ?????????? ??????????


Orodromeus makelai

0000000011 1110111000 0-010-0000 0010001000 --00010100 1????11001 1010001110 110-00-101 1111010122 10?0000?0? 0000100001 1111200111 10-1111102 111?001101 1101101000



Othnieliosaurus celer

?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ??????0122 ?01000000? ????100001 1101100111 10-1111102 1111001101 1111101?00


Pisanosaurus mertii

?????????? ????11???? ?????????? ?????????? ?????????? ?????11001 ?0?0??1??? ??0-00-?0? 1111?????? ?????????? ?????????? ?????????? ?????????? ??????000? 0????????0


Psittacosaurus

1110-01100 11--11100- 10100-0001 0010020011 0100001101 11-001?011 11P00015-- 1111110101 1111010022 1000000000 0000100001 1111200011 1111110102 111101?101 100?111000


Scelidosaurus harrisonii

0000000000 1100111120 0-00101001 0000010000 --00000100 ??0??11101 0011001010 110-00-101 1111000?11 000?00000? ????101000 0000000011 10-100-110 0101101101 1001101001


Scutellosaurus lawleri

?0?000000? ?10010???? 0-001000?? ?????110?? ????0????? 100?01110? ?0?100?0?? 110-00-101 11110?0111 ?????0000? 0?00101000 000?101001 10-10??100 0101001101 ??1???1??1


Stenopelix valdensis

?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ????????2? ?????????? ????101001 1101110011 11?111?11? 111????1?? ??0?111000



Stormbergia dangershoeki

?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ??????0?11 0????????? ????100000 000010P111 10-100-101 0101001101 1?111????0


Wannanosaurus yansiensis

?????????? ??????11?? 0-00110100 -0?????011 00111?0??? ?????0000? 10100?1??? 1111?101?? 01110????? ?????1011? ?????????? ?????????? ????????12 ????11111? ?????????0



Yinlong downsi

1110001001 1110111000 1010110000 00100?0010 01110???11 10100?1001 1000110311 210-0?-10? 11110????? ?????0???? ????100001 11??1??0?1 1111?1?1?? ???1?????? ?????????0



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