Aee 4

Journal of Systematic Palaeontology 6 (1): 1–40 Issued 22 February 2008
doi:10.1017/S1477201907002271 Printed in the United Kingdom

C The Natural History Museum
The phylogeny of the ornithischian

dinosaurs
Richard J. Butler∗
Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge, CB2 3EQ, UK &
Department of Palaeontology, The Natural History Museum, Cromwell Road, London, SW7 5BD, UK
Paul Upchurch†
Department of Earth Sciences, University College London, Gower Street, London, WC1E 6BT, UK
David B. Norman‡
Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge, CB2 3EQ, UK
SYNOPSIS Ornithischia is a familiar and diverse clade of dinosaurs whose global phylogeny has
remained largely unaltered since early cladistic analyses in the mid 1980s. Current understanding of
ornithischian evolution is hampered by a paucity of explicitly numerical phylogenetic analyses that
consider the entire clade. As a result, it is difficult to assess the robustness of current phylogenetic
hypotheses for Ornithischia and the effect that the addition of new taxa or characters is likely to have
on the overall topology of the clade.
The new phylogenetic analysis presented here incorporates a range of new basal taxa and charac-
ters in an attempt to rigorously test global ornithischian phylogeny. Parsimony analysis is carried out
with 46 taxa and 221 characters. Although the strict component consensus tree shows poor resolution
in a number of areas, application of reduced consensus methods provides a well-resolved picture
of ornithischian interrelationships. Surprisingly, Heterodontosauridae is placed as the most basal
group of all well-known ornithischians, phylogenetically distant from a stem-defined Ornithopoda,
creating a topology that is more congruent with the known ornithischian stratigraphical record. There
is no evidence for a monophyletic ‘Fabrosauridae’, and Lesothosaurus (the best-known ‘fabrosaur’)
occupies an unusual position as the most basal member of Thyreophora. Other relationships within
Thyreophora remain largely stable. The primitive thyreophoran Scelidosaurus is the sister taxon of
Eurypoda (stegosaurs and ankylosaurs), rather than a basal ankylosaur as implied by some previous
studies.
The taxonomic content of Ornithopoda differs significantly from previous analyses and basal
relationships within the clade are weakly supported, requiring further investigation. ‘Hypsilopho-
dontidae’ is paraphyletic, with some taxa (Agilisaurus, Hexinlusaurus, Othnielia) placed outside
of Ornithopoda as non-cerapodans. Ceratopsia and Pachycephalosauria are monophyletic and are
united as Marginocephalia; however, the stability of these clades is reduced by a number of poorly
preserved basal taxa.
This analysis reaffirms much of the currently accepted ornithischian topology. Nevertheless, in-
stability in the position and content of several clades (notably Heterodontosauridae and Ornithopoda)
indicates that considerable future work on ornithischian phylogeny is required and causes problems
for several current phylogenetic definitions.
KEY WORDS vertebrate palaeontology, Ornithischia, systematics, cladistics, Dinosauria
Contents
Introduction 1
Institutional abbreviations 2
Previous analyses of ornithischian phylogeny 2
Traditional classifications 2
Cladistic studies 3
Material and Methods 5
The aim of the analysis 5
Phylogenetic framework 5
Selection of ingroup taxa 5
∗ E-mail: [email protected]; †Email: [email protected]; ‡Email: [email protected]

2 R. J. Butler et al.
Supraspecific taxa 5
Included species level taxa 7
Excluded species level taxa 11
Selection of outgroup taxa 11
Analyses 12
Search methods 12
Testing the support for relationships 12
Randomisation tests 12
Bremer support 12
Bootstrapping 14
Results 16
Ornithischian monophyly 16
Pisanosaurus mertii 17
Heterodontosauridae 17
‘Fabrosaurids’ 18
Thyreophora 19
Basal neornithischians 19
Ornithopoda 20
Hypsilophodontidae 20
Marginocephalia 21
Ceratopsia 21
Pachycephalosauria 22
Conclusions 22
Stability, instability, and future directions in ornithischian phylogeny 22
Implications for phylogenetic taxonomy of Ornithischia 22
Acknowledgements 23
References 23
Appendix 1 – Specimens and references used for coding operational taxonomic units 29
Appendix 2 – Character list 30
Appendix 3 – Data matrix 35
Appendix 4 – Tree descriptions 40
Introduction tebrate faunas are dominated by saurischians (Weishampel

et al. 2004b). Ornithischians became much more abundant
during the Late Jurassic and Early Cretaceous and ornith-
The clade Ornithischia represents a major grouping within ischian diversity peaked during the Campanian stage of the
Dinosauria, the most familiar and widely popularised of Late Cretaceous. Late Triassic–Early Jurassic ornithischians
all extinct organisms. An ornithischian, Iguanodon Mantell, were generally small-bodied (1–2 m in length) bipedal curs-
1825, was the second dinosaur genus to be named, while ors (e.g. Lesothosaurus diagnosticus: Thulborn 1972; Sereno
of the three taxa explicitly included within Dinosauria by 1991a), but during the rest of the Mesozoic they diversified
Owen (1842), two (Iguanodon and Hylaeosaurus) were later into a considerable range of morphologies and sizes, with
recognised as ornithischians. Since these early discoveries, a many groups reverting to quadrupedality. The vast majority
large number of genera and species of ornithischians have of ornithischians are believed to have been herbivorous, al-
been named: a recent review (Weishampel et al. 2004a) though some basal forms have been interpreted as potentially
recognised over 180 valid genera. The earliest ornithischi- omnivorous (Barrett 2000).
ans are known from the Carnian stage of the Late Trias- One major problem in understanding ornithischian
sic (Casamiquela 1967) and the clade disappeared in the evolution is that, to date, there are few published numer-
mass extinctions at the end of the Cretaceous. During this ical phylogenetic analyses dedicated solely to Ornithischia
time ornithischians achieved a global distribution and are (Weishampel 2004). It is difficult, therefore, to assess the re-
now known from every continent, including Antarctica (e.g. lative phylogenetic support for ornithischian clades, to look
Hooker et al. 1991; Weishampel et al. 2004b). at the effects upon phylogenetic results of adding or deleting
Ornithischians appear to have been extremely scarce taxa or characters, or to test alternative hypotheses of taxon
during the Late Triassic (Sereno 1997) and remained un- or character evolution. Here, we present a new phylogen-
common (although apparently more diverse) during the Early etic analysis of Ornithischia, as part of an on-going study of
and Middle Jurassic; during this time interval terrestrial ver- ornithischian phylogeny.
Phylogeny of ornithischian dinosaurs 3
Institutional abbreviations ZDM = Zigong Dinosaur Museum, Dashanpu,

People’s Republic of China
BMNH = Natural History Museum, London, UK ZPAL = Institute of Paleobiology of the Polish
BRSMG = Bristol City Museum and Art Gallery, Bris- Academy of Sciences, Warsaw, Poland.
tol, UK
BP = Bernard Price Institute for Palaeontological
Research, Johannesburg, South Africa
BYU = Earth Science Museum, Brigham Young Previous analyses of
University, Provo, Utah, USA
CAMSM = Sedgwick Museum, University of Cam-
ornithischian phylogeny
bridge, Cambridge, UK
CEUM = Prehistoric Museum, College of Eastern
Traditional classifications
Utah, Price, Utah, USA Although material from the clade began to be described and
CV = Chongqing Natural History Museum, named in the early 19th century (Mantell 1822, 1825, 1833),
Chongqing, People’s Republic of China the recognition that ornithischians formed a grouping dis-
GCC = Museum of the Chendgu University of tinct from other reptiles (including other dinosaurs) was not
Technology (formerly Chendgu College of reached until the work of Seeley (1887). Seeley was the first
Geology), Chengdu, People’s Republic of to identify and articulate a fundamental morphological dicho-
China tomy within the then described dinosaurian taxa. He recog-
GI = Geological Institute, Ulaanbaatar, Mongo- nised two orders, distinguished mainly on the basis of pelvic
lia anatomy, that he named Ornithischia and Saurischia. Major
GSM = Geological Survey Museum, Keyworth, subdivisions of Seeley’s Ornithischia (Stegosauria, Ornitho-
UK poda, Ceratopsia) were identified by Marsh (1877a, 1881,
GZG = Geowissenschaftliches Zentrum der Uni- 1890) and Marsh (1894) later grouped these subdivisions
versität Göttingen, Göttingen, Germany together as the order Predentata, generally considered syn-
IGCAGS = Institute of Geology, Chinese Academy onymous with Ornithischia.
of Sciences, Beijing, People’s Republic of Nopcsa (1915) proposed a subdivision within Ornithis-
China chia, between the bipedal, unarmoured forms (ornithopods)
IVPP = Institute of Vertebrate Paleontology and and a new suborder that he named Thyreophora (compris-
Paleoanthropology, Bejing ing ankylosaurs, stegosaurs and ceratopsians). The group-
MCZ = Museum of Comparative Zoology, Harvard, ing Thyreophora was often ignored by later workers, but
USA the name was revived by Norman (1984) and Sereno (1984,
MB = Museum für Naturkunde, Berlin, Germany 1986), although its current usage (for a clade consisting of
MCF-PVPH = Museo Carmen Funes, Paleontologı́a de all armoured ornithischians) differs somewhat from that pro-
Vertebrados Plaza Huincul, Argentina posed by Nopcsa (i.e. in current usage ceratopsians are not
MNA = Museum of Northern Arizona, Flagstaff, members of Thyreophora).
USA Romer (1956) divided Ornithischia into four suborders:
MOR = Museum of the Rockies, Bozeman, Ornithopoda (including all bipedal forms), Stegosauria, An-
Montana, USA kylosauria and Ceratopsia. Most bipedal and relatively ‘un-
MPM = Museo Padre Molina, Rio Gallegos, Santa specialised’ taxa were included within Ornithopoda; other
Cruz, Argentina suborders were believed to be derived from within the or-
MWC = Museum of Western Colorado, Grand Junc- nithopods, although only the ceratopsians were explicitly
tion, Colorado, USA linked with a particular group of ornithopods (the psit-
OUM = University Museum of Natural History, Ox- tacosaurids).
ford, UK Thulborn (1971) suggested that most of the major
PVL = Fundación Miguel Lillo, Universidad groups of ornithischians (e.g. iguanodontids, pachycephalo-
Nacional de Tucumán, San Miguel de Tu- saurs, ceratopsians) were descended from a Late Triassic–
cumán, Argentina Late Cretaceous ‘hypsilophodont plexus’, an implicitly para-
ROM = Royal Ontario Museum, Toronto, Ontario, phyletic grade of small, primitive, bipedal ornithopods. An-
Canada kylosaurs and stegosaurs were considered to occupy a prim-
SAM-PK = Iziko South African Museum, Cape Town, itive position outside of this plexus, sharing ancestors with
South Africa the earliest hypsilophodontids. The details of this classifica-
SDSM = South Dakota School of Mines, Rapid City, tion were questioned by Galton (1972), who removed Ech-
South Dakota, USA inodon becklesii and Fabrosaurus australis from the family
SGWG = Sektion Geologische Wissenschaften Gre- Hypsilophodontidae to form the family Fabrosauridae (see
ifswald, Ernst-Moritz Universität, Greif- also Galton 1978), which included those taxa he considered
swald, Germany to represent the most basal known ornithischians. Maryańska
UC = University of Chicago, Chicago, USA & Osmólska (1974) emphasised the morphological distinct-
UCMP = University of California Museum of Pale- ness of the pachycephalosaurs, previously included as the
ontology, Berkeley, USA family Pachycephalosauridae within Ornithopoda, and cre-
YPM = Peabody Museum of Natural History, ated a new suborder of Ornithischia, Pachycephalosauria.
Yale University, New Haven, Connecticut, In summary, pre-cladistic ornithischian classifications
USA tended to recognise either four (Romer 1956) or five
(Maryańska & Osmólska 1974) suborders within Ornithis- dontosaurids were positioned as basal ornithopods, while
chia. One of these suborders (Ornithopoda) was an implicitly the clade containing Pachycephalosauria and Ceratopsia was
paraphyletic grouping of taxa, defined on the basis of ple- named Marginocephalia. Sereno followed Norman (1984) in
siomorphic characters (e.g. bipedality). Only a few workers restricting Thyreophora to ankylosaurs, stegosaurs and two
(e.g. Thulborn 1971; Galton 1972) attempted to identify the basal armoured forms (Scutellosaurus lawleri, Scelidosaurus
pattern of interrelationships between or within these clades. harrisonii), while thyreophorans and cerapodans were united
as the clade Genasauria. Sereno continued to consider Fab-
rosauridae polyphyletic and positioned Lesothosaurus dia-
Cladistic studies gnosticus as the sister taxon of Genasauria.
The first cladistic studies of ornithischian phylogeny were Following the work of Sereno (1986), ornithischian
published simultaneously by Norman (1984), Milner & workers tended to focus on relationships within the major
Norman (1984) and Sereno (1984); the results of Norman ornithischian clades; e.g. the phylogeny of basal Ornitho-
(1984) and Sereno (1984) are shown in Fig. 1A, B. Norman poda has been analysed by Weishampel & Heinrich (1992),
(1984) proposed that Ornithischia could be divided into two Winkler et al. (1997), Scheetz (1998, 1999), Weishampel
major groupings: Thyreophora, comprising the ankylosaurs et al. (2003) and Norman et al. (2004c), amongst others.
and stegosaurs, and Ornithopoda, which Norman expanded to However, there have been only a few attempts to test the
include ceratopsians. Norman positioned fabrosaurs as basal global phylogeny of Ornithischia.
ornithopods, suggested that ceratopsians and iguanodontians Sereno (1997: figs 1, 2) presented an ornithischian
(referred to as ‘dryosauroideans’ in his cladogram) shared cladogram, but this was not supported by a data matrix or in-
a common ancestor to the exclusion of hypsilophodontids, formation about analyses. In a review paper, Sereno (1999a)
but considered the position of heterodontosaurids and pachy- considered ornithischians within a larger scale analysis of di-
cephalosaurs to be problematic and unresolved. The phylo- nosaurian phylogeny. This included nine separate data sets,
geny of Milner & Norman (1984) concentrated on relation- each of which analysed separate portions of the dinosaur-
ships within Ornithopoda and effectively represented a subset ian tree. Four of these data sets (data sets 1–4) dealt with
of the analysis of Norman (1984). ornithischians. Data set 1 analysed basal dinosaurian phylo-
The phylogeny presented by Sereno (1984) differed geny and, within this framework, tested both ornithischian
significantly from that of Norman (1984). Sereno proposed monophyly and interrelationships. Data sets 2–4 analysed the
that Fabrosauridae was polyphyletic and positioned Lesotho- within-clade phylogeny of Thyreophora, Ornithopoda and
saurus diagnosticus (previously included within Fabrosaur- Marginocephalia. Results generally supported the findings of
idae) as the most basal known ornithischian. His conception Sereno (1986) and differed only in the inclusion of additional
of Ornithopoda was much more restrictive than that of previ- taxa (e.g. the basal thyreophoran Emausaurus ernsti) and the
ous workers and excluded ceratopsians, pachycephalosaurs identification of the Late Triassic taxon Pisanosaurus mertii
and ‘fabrosaurs’, while including heterodontosaurids. His as the most basal ornithischian. Monophyly of Thyreophora,
conception of Thyreophora also differed substantially from Ornithopoda and Marginocephalia (and a taxonomic content
that of Norman (1984), comprising a clade consisting of an- for these clades consistent with the phylogenetic results of
kylosaurs, stegosaurs, pachycephalosaurs and ceratopsians. Sereno (1986)), was assumed prior to analysis.
The phylogeny of Cooper (1985) was similar in many Buchholz (2002) carried out an ornithischian analysis
aspects (Fig. 1C) to that of Sereno (1984). However, Cooper with 27 taxa and 97 characters. Taxonomic sampling was re-
positioned Heterodontosauridae as the sister group to the stricted mostly to taxa generally considered as ornithopods,
Pachycephalosauria–Ceratopsia clade and considered Fab- with Marginocephalia coded as a composite taxon and basal
rosauridae to represent the most basal clade within Ornitho- thyreophorans excluded from the analysis. Buchholz found
poda. support for a sister-grouping of heterodontosaurids and mar-
Maryańska & Osmólska (1985) outlined a phylogeny ginocephalians and for paraphyly of hypsilophodontids. Un-
(Fig. 1D) that differed in several key points from that of fortunately, although a character-list was published, a matrix
Norman (1984) and Sereno (1984). Maryańska & Osmólska was not provided and these results cannot be reassessed.
proposed that ankylosaurs and stegosaurs did not share a Liu (2004) tested the global phylogeny of Ornithischia
common ancestor but, instead, represented serial outgroups with a large-scale analysis including 44 taxa and 326 char-
to more derived ornithischians, and that heterodontosaurids acters. Interesting results were reported: Fabrosauridae and
formed the sister group to a clade consisting of Ornitho- Hypsilophodontidae were found to be paraphyletic, Leso-
poda (including Lesothosaurus diagnosticus), Pachycephalo- thosaurus was positioned as a basal member of Thyreo-
sauria and Ceratopsia. Following Sereno (1984), Maryańska phora, Agilisaurus louderbacki (often considered to repres-
& Osmólska united Pachycephalosauria and Ceratopsia to ent a basal ornithopod, e.g. Norman et al. 2004c) grouped
form a clade to the exclusion of ornithopods. outside of Cerapoda and Marginocephalia and Iguanodontia
The most influential published work on ornithischian were united as sister groups. Unfortunately, this analysis has,
phylogeny was produced by Sereno (1986) and his res- to date, been published in abstract form only. It is, therefore,
ults (Fig. 1E) have dominated subsequent understanding impossible, at this stage, to reanalyse the data. However, the
of ornithischian phylogeny (see, e.g. Weishampel et al. reported results of Liu (2004) clearly highlight the import-
1990, 1992, 2004a; Fastovsky & Weishampel 1996, 2005; ance of new phylogenetic analyses.
Currie & Padian 1997; Sereno 1997, 1999a). Sereno (1986) One of us (Butler 2005) included a cladistic analysis
modified his earlier (Sereno 1984) hypothesis by uniting or- (23 taxa, 73 characters) of Ornithischia within a review of the
nithopods, pachycephalosaurs and ceratopsians (following ‘fabrosaurid’ ornithischians of the upper Elliot Formation of
Norman (1984) and Maryańska & Osmólska (1985)) in a southern Africa. That analysis recovered interesting results,
clade that he termed Cerapoda. Within Cerapoda, hetero- including positioning heterodontosaurids and the Middle
Figure 1 Previous numerical ornithischian phylogenies, simplified and redrawn from the original publications. A, Norman (1984); B, Sereno
(1984); C, Cooper (1985); D, Maryańska & Osmólska (1985); E, Sereno (1986 ). H1, H2, alternative positions proposed by Norman (1984) for the
clade Heterodontosauridae; P1, P2, alternative positions proposed by Norman (1984) for the clade Pachycephalosauria.
Jurassic taxa Agilisaurus louderbacki and Hexinlusaurus Table 1 provides phylogenetic definitions for ornithis-
multidens outside of Cerapoda; that analysis represented an chian clade names discussed in the text (modified from Ser-
early iteration of the analysis presented here and is super- eno 1998, 1999b; Wagner 2004). There is a conflict in the lit-
seded by the present study. erature between the names Neornithischia (Cooper 1985) and
Although a number of major phylogenetic studies of Or- Cerapoda (Sereno 1986), which have both been applied to
nithischia have been published (Norman 1984; Sereno 1984, the clade consisting of ornithopods, pachycephalosaurs and
1986, 1999a; Cooper 1985; Maryańska & Osmólska 1985), ceratopsians (e.g. Sereno 1999a; Weishampel 2004). Here,
the majority have failed to include crucial information, in- we follow Buchholz (2002) and Barrett et al. (2005) by us-
cluding: (1) a character–taxon data matrix; (2) details of the ing both names: Neornithischia is applied to a stem-based
specimens and references used in coding operational taxo- clade, while Cerapoda is used for a node-based clade. In ad-
nomic units; (3) tree searching methods; (4) the number and dition, our use of Ornithopoda differs from that of those au-
scores of the most parsimonious trees recovered by search thors who have defined this taxon as a node-based clade util-
methods; (5) tests of data robustness and support for particu- ising Heterodontosaurus as an internal specifier (e.g. Sereno
lar clades (e.g. bootstrapping, decay analysis). Most previous 1998); we instead use Ornithopoda for a stem-based clade
studies have simply presented a fully-resolved tree and lists (Buchholz 2002; Wagner 2004; Norman et al. 2004c).
of apomorphies for particular clades. It is neither possible to
independently rerun these analyses and recover their results, Selection of ingroup taxa
nor is it possible to assess data robustness, the relative sup-
port for clades, or the support for alternative phylogenetic A taxonomic review of Ornithischia was carried out and a
hypotheses. number of supraspecific, species level and outgroup oper-
With the exception of the preliminary study of Butler ational taxonomic units (OTUs) were selected. Weishampel
(2005), Sereno (1999a) is the only published ornithischian et al. (1990, 1992, 2004a) served as the source for this review.
analysis that includes a data matrix that can be rerun and Coding of taxa for cladistic analysis was based, where pos-
reanalysed. However, there are problems with this analysis. sible, on first-hand examination of specimens, supplemented
Only a limited number of ornithischian taxa were included with information from the literature. Appendix 1 provides de-
and, in addition, monophyly of major clades (such as Mar- tails of the references and specimens used for coding OTUs.
ginocephalia) was assumed prior to analysis; unfortunately, The choice of ingroup taxa is discussed in greater detail be-
monophyly of clades such as Marginocephalia are questions low and outgroup taxa are discussed in the following section.
that ornithischian analyses still need to resolve.
Supraspecific taxa
A number of authors (e.g. Wiens 1998; Prendini 2001) have
suggested that supraspecific taxa should be avoided in phylo-
Material and Methods genetic analysis when possible, as the coding of such taxa
is problematic, and simulations tend to suggest that splitting
The aim of the analysis such taxa into species level terminals provides better results.
Ideally, therefore, any analysis of Ornithischia should util-
The aim of this analysis is to test the global phylogeny of Or- ise only species level terminal taxa. Nevertheless, the use of
nithischia, concentrating on the phylogenetic relationships species level taxa as exemplars for major clades was con-
of basal forms. Questions concerning the monophyly of the sidered impractical for this analysis for a number of reasons.
Dinosauria, the phylogenetic relationships of basal ‘dino- First, the choice of exemplar taxa is not always obvious. For
sauriformes’ and basal saurischians, and the relationships of instance, the clade Ankylosauria is nearly universally accep-
derived taxa within the major ornithischian clades, are bey- ted as monophyletic and is well-supported by anatomical
ond the scope of this analysis. The intention is to test the evidence. However, there is little consensus as to phylogen-
phylogenetic framework upon which the current understand- etic relationships within Ankylosauria (e.g. Kirkland 1998;
ing of ornithischian evolution is based. Carpenter 2001; Vickaryous et al. 2001, 2004; Parish 2003)
and justifying the use of one taxon, or several taxa, as exem-
plars is difficult. In addition, many of the apparently basal an-
Phylogenetic framework kylosaur taxa that might be used as exemplars are fragment-
All published cladistic analyses have shown Dinosauria to ary (e.g. Cedarpelta bilbeyhallorum, Mymoorapelta maysi),
form a monophyletic clade (e.g. Gauthier 1986; Benton incompletely described in the literature (Gastonia burgei), or
& Clark 1988; Novas 1996; Sereno 1999a; Benton 2004; based upon juvenile material (Liaoningosaurus paradoxus).
Langer 2004; Langer & Benton 2006). Dinosauria includes In such a situation it can be advantageous to code a supra-
two subclades: Saurischia and Ornithischia (Gauthier 1986; specific taxon to represent the clade. The use of supraspecific
Sereno 1986). A number of outgroups to Dinosauria have taxa has the additional advantage of reducing the number of
been identified; the most proximate of which appear to be sev- OTUs that must be included in the analysis. This is important,
eral ‘dinosauriform’ taxa, exemplified by Lagerpeton (Ser- because it allows heuristic searches to be carried out in an
eno & Arcucci 1993), Marasuchus (Sereno & Arcucci 1994) acceptable timeframe and allows much more detailed exam-
and Silesaurus (Dzik 2003; Langer & Benton 2006). Suc- ination of the data. For these reasons, selected supraspecific
cessively more distant outgroups to Dinosauria within Archo- taxa were included in this analysis.
sauria include Pterosauria (although see Bennett 1996; Peters As outlined by Bininda-Emonds et al. (1998), the cor-
2000), Scleromochlus, Crurotarsi, Proterochampsidae, Eu- rect use of supraspecific taxa in phylogenetic analyses has
parkeria, Erythrosuchus and Proterosuchus (Sereno 1991b; two requirements. Firstly, the taxa must be monophyletic.
Benton 2004). Only supraspecific taxa that are generally accepted as
Table 1 Phylogenetic definitions for the major ornithischian clades discussed in this analysis (modified from: Sereno 1998, 1999b; Buchholz
2002; Wagner 2004).
Clade name Phylogenetic definition

Dinosauria Owen, 1842 Triceratops horridus Marsh, 1889, Passer domesticus (Linnaeus, 1758), their most
recent common ancestor and all descendents.
Saurischia Seeley, 1887 All dinosaurs more closely related to Passer domesticus (Linnaeus, 1758) than to
Triceratops horridus Marsh, 1889.
Ornithischia Seeley, 1887 All dinosaurs more closely related to Triceratops horridus Marsh, 1889 than to either
Passer domesticus (Linnaeus, 1758), or Saltasaurus loricatus Bonaparte & Powell,
1980.
Genasauria Sereno, 1986 Ankylosaurus magniventris Brown 1908, Stegosaurus stenops Marsh, 1877a,
Parasaurolophus walkeri Parks, 1922, Triceratops horridus Marsh, 1889,
Pachycephalosaurus wyomingensis (Gilmore, 1931), their most recent common
ancestor and all descendents.
Thyreophora Nopcsa, 1915 All genasaurians more closely related to Ankylosaurus magniventris Brown, 1908
than to Parasaurolophus walkeri Parks, 1922, Triceratops horridus Marsh, 1889, or
Pachycephalosaurus wyomingensis (Gilmore, 1931).
Eurypoda Sereno, 1986 Ankylosaurus magniventris Brown, 1908, Stegosaurus stenops Marsh, 1877a, their
most recent common ancestor and all descendents.
Ankylosauria Osborn, 1923 All ornithischians more closely related to Ankylosaurus magniventris Brown, 1908
than to Stegosaurus stenops Marsh, 1877a.
Stegosauria Marsh, 1877a All ornithischians more closely related to Stegosaurus stenops Marsh, 1877a than to
Ankylosaurus magniventris Brown, 1908.
Neornithischia Cooper, 1985 All genasaurians more closely related to Parasaurolophus walkeri Parks, 1922, than
to Ankylosaurus magniventris Brown, 1908 or Stegosaurus stenops Marsh, 1877a.
Cerapoda Sereno, 1986 Parasaurolophus walkeri Parks, 1922, Triceratops horridus Marsh, 1889, their most
recent common ancestor and all descendents.
Ornithopoda Marsh, 1881 All genasaurians more closely related to Parasaurolophus walkeri Parks, 1922, than
to Triceratops horridus Marsh, 1889
Marginocephalia Sereno, 1986 Triceratops horridus Marsh, 1889, Pachycephalosaurus wyomingensis (Gilmore,
1931), their most recent common ancestor and all descendents.
Ceratopsia Marsh, 1890 All marginocephalians more closely related to Triceratops horridus Marsh, 1889 than
to Pachycephalosaurus wyomingensis (Gilmore, 1931).
Pachycephalosauria Maryańska & Osmólska, 1974 All marginocephalians more closely related to Pachycephalosaurus wyomingensis
(Gilmore, 1931) than to Triceratops horridus Marsh, 1889.
monophyletic were utilised here. Secondly, it must be pos- (1998), Vickaryous et al. (2001), Hill et al. (2003) and
sible to code them as a single OTU in a manner that maintains Vickaryous et al. (2004). The phylogenetic analysis of
their position on a cladogram with respect to a solution in- Carpenter (2001) is not used, because it utilised a com-
cluding all species. Several authors (e.g. Bininda-Emonds partmentalisation technique and does not, therefore, rep-
et al. 1998; Wiens 1998) suggest that the ‘ancestral’ method, resent a global phylogeny.
whereby the character states of a hypothetical ancestor (the 2. Stegosauria. Stegosauria is defined as all taxa more
‘groundplan’) are reconstructed on the basis of prior phylo- closely related to Stegosaurus stenops than to Ankylo-
genetic analyses, is the most successful method of coding saurus magniventris (Sereno 1998). A full listing of in-
supraspecific taxa. The ‘ancestral’ method (the methodology cluded taxa and synapomorphies supporting monophyly
is outlined by Langer & Benton 2006) was used in this ana- of the clade is given in Galton & Upchurch (2004b). Ste-
lysis. The eight supraspecific OTUs used are discussed in gosaurs form a relatively small but well-known and well-
greater detail below: supported clade of ornithischians, known mostly from the
Middle to Late Jurassic, with fragmentary forms known
1. Ankylosauria. Ankylosauria is defined as all taxa more from the Early Cretaceous.
closely related to Ankylosaurus magniventris than to Sereno & Dong (1992) provided the first phylo-
Stegosaurus stenops (Sereno 1998) and includes the genetic analysis of stegosaurs, but considered only a
subclades Ankylosauridae and Nodosauridae. Included few taxa. They proposed that Huayangosaurus taibaii
taxa and diagnostic features are listed by Vickaryous represents the most basal member of the clade, with
et al. (2004). The known temporal range of the clade is Dacentrurus armatus positioned as the sister-group
Callovian to Maastrichtian (Middle Jurassic–Late Creta- to all more derived stegosaurs. Galton & Upchurch
ceous). (2004b) have provided the most complete analysis to
The most recent review of Ankylosauria (Vickaryous date; the basal positions of Huayangosaurus and Da-
et al. 2004) recognised over 40 valid species. The phylo- centrurus were confirmed by their analysis, but little
geny assumed here for character coding represents a con- resolution was found amongst more derived stego-
sensus of the following published phylogenies: Kirkland saurs.
3. Rhabdodontidae. Rhabdodontidae is defined as Zalmoxes taxa ranging from the Turonian to the Maastrichtian (Late
robustus, Rhabdodon priscus, their common ancestor and Cretaceous). Synapomorphies of this node are given by
all of its descendents (Weishampel et al. 2003) and the Sereno (2000), Makovicky (2001) and Makovicky &
temporal range of the clade extends from the Late San- Norell (2006).
tonian to the Maastrichtian (Late Cretaceous). Synapo- Neoceratopsian phylogeny has undergone a number of
morphies supporting the clade are given by Weishampel rigorous studies in recent years (Sereno 2000; Makovicky
et al. (2003). 2001; Xu et al. 2002; You & Dodson 2003, 2004;
4. Dryosauridae. Dryosauridae includes the taxa Dry- Chinnery 2004; Makovicky & Norell 2006) and a broad-
osaurus altus, Dryosaurus lettowvorbecki, Valdosaurus scale consensus has been reached that Liaoceratops and
canaliculatus and Valdosaurus nigeriensis (Norman Archaeoceratops represent basal taxa and that other neo-
2004) and is defined as Dryosaurus altus and all taxa ceratopsian taxa form a distinct clade. This clade of
more closely related to it than to Parasaurolophus walkeri more derived neoceratopsians remains unnamed and,
(Sereno 1998). The clade is known from the Late Jurassic generally, comprises a clade known as Leptoceratop-
and Early Cretaceous. Potential synapomorphies of this sidae and a clade known as Coronosauria, which in turn
clade include: lacrimal inserts into notch in the maxilla; comprises Protoceratopsidae and Ceratopsoidea (Sereno
very wide brevis shelf on the ilium; large, deep pit on the 2000; Makovicky 2001; Xu et al. 2002; Chinnery 2004).
femoral shaft, at the base of the fourth trochanter; digit I
of the pes is absent or vestigal.
5. Ankylopollexia. Ankylopollexia is defined as Campto- Included species level taxa
saurus dispar, Parasaurolophus walkeri, their common Following the selection and definition of supraspecific taxa,
ancestor and all descendents (Sereno 1998). A listing of the status of all taxa not included in one of these derived
included taxa and synapomorphies supporting monophyly clades was assessed, using first-hand observations and the
of the clade is given by Norman (2004). The clade ex- literature. For each taxon a decision was made as to whether
tends from the Kimmeridgian to the Maastrichtian (Late it should be included in phylogenetic analysis or not. Included
Jurassic–Late Cretaceous). species level taxa are discussed in this section; excluded spe-
Ankylopollexia was erected by Sereno (1986) for or- cies level taxa (and the reasons for exclusion) are discussed
nithopods exhibiting derived features of the teeth and in the following section.
manus, in particular the modification of manual digit I to Abrictosaurus consors (Thulborn, 1974) is known from
accommodate a spine-like pollex. Within this clade are a single partial skull and postcranial skeleton (BMNH
Camptosauridae and Styracosterna, both of which have RUB54, holotype; formerly UCL B54) from the upper
been given stem-based phylogenetic definitions by Sereno Elliot Formation of Lesotho (Early Jurassic: Hettangian–
(1998). The monophyly of Ankylopollexia is universally Sinemurian). Autapomorphies have not previously been
supported by phylogenetic analysis and its interrelation- defined for Abrictosaurus and this taxon is provisionally dia-
ships are relatively well understood (e.g. Norman 2002, gnosed by the following combination of characters: arched
2004). diastema between premaxilla and maxilla present; enlarged
6. Pachycephalosauridae. Pachycephalosauridae is defined caniniform teeth absent from the premaxilla and dentary. The
as all taxa more closely related to Pachycephalosaurus single specimen of Abrictosaurus was initially described as a
wyomingensis than to either Homalocephale calathocer- new species of the heterodontosaurid Lycorhinus (Thulborn
cos or Goyocephale lattimorei. Known taxa are restricted 1974); however, it can clearly be distinguished from Ly-
to the Late Cretaceous (Campanian–Maastrichtian) and corhinus on the basis of dental characters (Hopson 1975).
synapomorphies are given in Sereno (2000). Hopson (1975) tentatively referred the specimen BMNH
A number of explicit, numerical phylogenetic analyses A100 (discussed below) to Abrictosaurus; however, this re-
of Pachycephalosauridae have been carried out in recent ferral has not been supported by subsequent work. Abricto-
years (Sereno 2000; Williamson & Carr 2002; Sullivan saurus and BMNH A100 are included here as separate OTUs.
2003; Maryańska et al. 2004).These studies indicate a Agilisaurus louderbacki Peng, 1990 is known from a
basal position for Stegoceras and the existence of a de- complete, articulated skull and postcranial skeleton (ZDM
rived clade containing Tylocephale, Prenocephale and T6011, holotype) from the Lower Shaximiao Formation of
Pachycephalosaurus, amongst others. Sichuan Province, People’s Republic of China (Middle Jur-
7. Psittacosauridae. Psittacosauridae is defined as all taxa assic: ?Bajocian, Chen et al. 1982; ?Bathonian–Callovian,
more closely related to Psittacosaurus mongoliensis than Dong & Tang 1984). Agilisaurus can be distinguished
to Triceratops horridus and contains the genera Hong- by the following autapomorphies: presence of a palpeb-
shanosaurus and Psittacosaurus. Synapomorphies of the ral/supraorbital bar that traverses the width of the orbit;
clade are given by Sereno (2000). The genus Psit- anteriormost dentary teeth conical, resembling premaxillary
tacosaurus is probably the most speciose and diverse of teeth; presence of a series of low, anterolaterally directed
all dinosaur genera, although the exact number of spe- ridges on the orbital portion of the frontal; presence of an
cies recognised is controversial (Sereno 1990b; You & excavated area immediately anterior to the antorbital fossa
Dodson 2004). Relationships within the clade are poorly (modified from Barrett et al. 2005).
understood. Members of Psittacosauridae are known from Anasibetia saldiviai Coria & Calvo, 2002 is known
the Early Cretaceous. from a partial skeleton with skull fragments (MCF-PVPH-
8. Unnamed taxon (Coronosauria + Leptoceratopsidae). 74, holotype) as well as a number of referred speci-
This unnamed clade is defined as Leptoceratops gracilis, mens (see Coria & Calvo 2002) from the Lisandro Form-
Protoceratops andrewsi and Triceratops horridus, their ation of Neuquén Province, Argentina (Late Cretaceous:
common ancestor and all of its descendents and includes Cenomanian). This taxon can be diagnosed on the basis of its
anteroventrally orientated occipital condyle and the presence triangular plate-like palpebral, the robust lateral margin of
of an ilium with preacetabular process longer than 50% of which bears a number of low ridges.
the total ilium length (modified from Coria & Calvo 2002). Gasparinisaura cincosaltensis Coria & Salgado, 1996
Archaeoceratops oshimai Dong & Azuma, 1997 is is known from numerous specimens (see Coria & Salgado
known from a well-preserved skull and postcranial material 1996; Salgado et al. 1997) from the Rio Colorado Forma-
(IVPP V11114, holotype; IVPP V11115, paratype) from the tion of Patagonia, Argentina (Late Cretaceous: Coniacian–
Xinminbao Group of Gansu Province, China (Early Creta- Santonian) and is diagnosed by the following characters:
ceous: Aptian–Albian) and is characterised by the presence of anteroposteriorly wide ascending process of lacrimal con-
an excavation on the lateral surface of the ischiadic peduncle tacts ventral process of postorbital posteriorly; infratemporal
of the ilium, as well as by a unique character combination fenestra bordered entirely ventrally by quadratojugal; apex
(modified from You & Dodson 2003). of arched dorsal margin of infratemporal fenestra positioned
The specimen BMNH A100 (formerly UCL A100) posterior to mandibular articulation; fully fused greater and
comprises a partial, disarticulated skull from the upper El- lesser trochanters; condylid of femur laterally positioned
liot Formation of South Africa. Assignment of this speci- (modified from Coria & Salgado 1996).
men to Lycorhinus (Thulborn 1970b, 1974; Gow 1990) was Goyocephale lattimorei Perle et al. 1982 is known from
not based on unique characters, but on general similarity. A a relatively complete skeleton with partial skull (GI SPS
number of subsequent authors criticised the referral of this 100/1501, holotype), from an unnamed unit, Mongolia (Late
specimen to Lycorhinus: Galton (1973a: caption to fig. 2) Cretaceous: ?late Santonian or early Campanian). One auta-
referred BMNH A100 to Heterodontosaurus sp.; Charig & pomorphy has been identified: the lateral margin of the skull
Crompton (1974) considered BMNH A100 to be generic- is sinuous in dorsal view (Sereno 2000).
ally distinct from both Heterodontosaurus and Lycorhinus; Heterodontosaurus tucki Crompton & Charig, 1962 is
while Hopson (1975) provisionally referred BMNH A100 to known from a nearly complete skull (SAM-PK-K337, holo-
Abrictosaurus. type) from the Clarens Formation (= Cave Sandstone) of
The taxonomy of the Elliot Formation heterodontosaur- South Africa (Early Jurassic: Sinemurian) and a referred
ids is poorly resolved and requires further work. At present skull and postcranial skeleton (SAM-PK-K1332, Santa Luca
there is no consensus as to the taxonomic status of BMNH et al. 1976; Santa Luca 1980) from the upper Elliot Form-
A100; however, this specimen is known from relatively com- ation of South Africa. A number of features may be auta-
plete and informative cranial remains and has received a de- pomorphic for this taxon, although it is possible that some
tailed description (Thulborn 1970b) and is thus included in may prove to be present in other, poorly known, heterodon-
the phylogenetic analysis here as a separate OTU. tosaurids, or may eventually prove to be ornithischian ple-
Bugenasaura infernalis Galton, 1995 was erected for a siomorphies. These possible autapomorphies include: dorsal
partial skull and postcranial fragments (SDSM 7210, holo- process of premaxilla does not form contact with nasals; an-
type) from the Hell Creek Formation of South Dakota, USA terior, accessory opening present within the antorbital fossa;
(Late Cretaceous: Maastrichtian). This taxon is diagnosed by squamosal–quadratojugal contact is anteroposteriorly broad;
the following features: no edentulous region at the anterior paroccipital processes are very deep dorsoventrally; paired,
end of the premaxilla; very deeply recessed cheek tooth row, deep recesses on the ventral surface of the basisphenoid;
with a massive and deep dentary and a very prominent over- basisphenoid processes are extremely elongated; cingulum
hanging ridge (with a braided appearance) on the ventral is completely absent on cheek-teeth; ischium with elongate
part of the maxilla; distal end of palpebral obliquely trun- flange on lateral margin.
cated with ridges medially (modified from Galton 1999). A Hexinlusaurus multidens (He & Cai, 1983) is known
new skeleton of Bugenasaura is known (MOR 979; R.J.B. from two partial skulls and postcranial skeletons (ZDM
pers. obs. 2004); however, this specimen has not been fully T6001, holotype; ZDM T6002, paratype) from the Lower
prepared and is currently undescribed. Shaximiao Formation of China. Hexinlusaurus can be distin-
Chaoyangsaurus youngi Zhao et al. 1999 is based upon guished by the presence of a marked concavity that extends
a partial skull and fragmentary postcranial elements (IG- over the lateral surface of the postorbital (Barrett et al. 2005).
CAGS V371, holotype) from the Tuchengzi Formation of Homalocephale calathocercos Maryańska &
Liaoning Province, China (Middle or Late Jurassic: Middle Osmólska, 1974 is known only from a skull and par-
Jurassic, Zhao et al. 1999; Tithonian, Weishampel et al. tial postcranial skeleton (GI SPS 100/1201, holotype) from
2004b). Chaoyangsaurus is distinguished by the follow- the Nemegt Formation of Mongolia (Late Cretaceous: ?late
ing autapomorphic features: quadratojugal overlaps posterior Campanian or early Maastrichtian). Homalocephale is
side of the quadrate shaft; quadrate slopes strongly anteriorly; diagnosed by the presence of a postacetabular process of the
ridge present between the planar lateral and ventral surfaces ilium that is crescent-shaped and ventrally deflected (Sereno
of the angular (modified from Zhao et al. 1999). 2000).
Echinodon becklesii Owen, 1861b is based upon frag- Hypsilophodon foxii Huxley, 1869 is known from nu-
mentary cranial material (see Norman & Barrett 2002) from merous specimens (see Galton 1974a) from the Wessex
the Purbeck Formation of England (Early Cretaceous: Berri- Formation of the Isle of Wight, UK (Early Cretaceous: Bar-
asian). Echinodon can be diagnosed by the presence of one, remian). Autapomorphies have not been previously defined
or possibly two, caniniform teeth situated at the anterior end for Hypsilophodon, but include the presence of a large fora-
of the maxilla (Norman & Barrett 2002). men in the ascending process of the maxilla that communic-
Emausaurus ernsti Haubold, 1990 is known from a par- ates medially with the antorbital fossa (Galton 1974a: fig. 3).
tial skull and postcranial fragments (SGWG 85, holotype), Although material of Hypsilophodon has been reported from
from an unnamed unit in Germany (Early Jurassic: Toarcian). continental Europe (Sanz et al. 1983) and North America
Emausaurus can be diagnosed by the possession of a large, (Galton & Jensen 1979), none of this material can be
confidently referred to this taxon and, at present, Hypsilo- extremely fragmentary and many elements were erroneously
phodon is only known from the UK. identified in the original description. A full review and rede-
Jeholosaurus shangyuanensis Xu et al. 2000 is known scription is being prepared (R. J. B. & Q. Zhao, unpublished
from two specimens (IVPP V12529, holotype; IVPP results). Although Sereno (2000) has suggested that autapo-
V12530, referred) from the Yixian Formation of Liaoning morphic features are absent, the presence of prominent vent-
Province, China (Early Cretaceous) and is characterised by ral grooves on the most posterior dorsal vertebrae appears
the following combination of characters: six premaxillary to be autapomorphic for Micropachycephalosaurus and this
teeth; foramina present on dorsal surface of nasal; large fo- taxon is here included in the phylogenetic analysis.
ramen present in quadratojugal; predentary about 1.5 times Orodromeus makelai Horner & Weishampel, 1988 is
as long as the premaxilla; pedal phalanx 3–4 times longer known from abundant and well-preserved material (see
than other phalanges of pedal digit 3 (modified from Xu Scheetz 1999) from the Upper Two Medicine and Judith
et al. 2000). River formations of Montana, USA (Late Cretaceous: ?Late
Lesothosaurus diagnosticus Galton, 1978 is known Campanian). Orodromeus is characterised by the following
from a number of nearly complete skulls and disarticulated combination of characters: prominent boss on anterolateral
postcranial skeletons, while Stormbergia dangershoeki But- maxilla; subnarial depression on premaxilla–maxilla bound-
ler, 2005 is known from three partial skeletons. Both taxa are ary; midline depression on nasals; boss on jugal; postorbital
from the upper Elliot Formation of South Africa and Leso- with distinct projection into orbit; dentition plesiomorphic
tho. A full discussion of the hypodigm and diagnosis of each with ridges absent lingually and labially.
taxon can be found in Butler (2005). Othnielia rex (Marsh, 1877b) is known from a num-
Liaoceratops yanzigouensis Xu et al. 2002 is known ber of specimens (YPM 1915, holotype, left femur; referred
from two complete skulls (IVPP V12738, holotype; IVPP specimens (see Galton 1983) include: BYU ESM-163R, ar-
V12633, referred specimen) from the Lower Yixian Forma- ticulated, near-complete postcranial skeleton described by
tion of Liaoning Province, China (Early Cretaceous). Liao- Galton & Jensen 1973) from the Morrison Formation of
ceratops is characterised by the following features: sutures the USA (Late Jurasssic: Kimmeridgian–Tithonian). YPM
between premaxilla, maxilla, nasal and prefrontal intersect- 1915 is the holotype of Nanosaurus rex Marsh, 1877b, which
ing at a common point high on the side of the snout; posses- was made the type species of the genus Othnielia by Galton
sion of several tubercles on the ventral margin of the angular; (1977). This specimen (an isolated femur) lacks obvious auta-
a foramen on the posterior face of the quadrate near the artic- pomorphies, although it may be diagnosable on the basis of a
ulation with the quadratojugal; small tubercle on the dorsal unique character combination. Referral of specimens to Oth-
border of the foramen magnum; thick posterior border of the nielia follows Galton (1983), pending a review of the validity
parietal frill (Xu et al. 2002). of this taxon.
Lycorhinus angustidens Haughton, 1924 is known from Parksosaurus warreni (Parks, 1926) is known from a
a left dentary (SAM-PK-K3606, holotype) and two provi- single, relatively complete, skull and skeleton (ROM 804,
sionally referred specimens (BP/1/4244, left maxilla, holo- holotype) from the Horseshoe Canyon Formation of Alberta,
type of Lanasaurus scalpridens Gow, 1975, referred to Ly- Canada (Late Cretaceous: Maastrichtian). Parksosaurus is
corhinus angustidens by Gow 1990; BP/1/5253, left max- distinguished by a dorsoventrally broad posterolateral pro-
illa, referred to Lycorhinus angustidens by Gow 1990) from cess of the premaxilla and a postorbital process of the jugal
the upper Elliot Formation of South Africa. There has been that expands posterodorsally.
considerable controversy over the validity of Lycorhinus an- Pisanosaurus mertii Casamiquela, 1967 is known from
gustidens. Haughton (1924) named Lycorhinus for a partial a single partial skeleton (PVL 2577, holotype) from the Is-
left dentary that he believed represented a cynodontid syn- chigualasto Formation (Late Triassic: Carnian) of Argentina.
apsid. Crompton & Charig (1962) reidentified Lycorhinus as As discussed by Sereno (1991a), this taxon can be diagnosed
a heterodontosaurid and, later Charig & Crompton (1974) by the following characters: anteroposterior depth of distal
considered it a nomen dubium. Thulborn (1970b) assigned end of the tibia is greater than maximum transverse width;
the specimen BMNH A100 to Lycorhinus; however, this as- calcaneum is transversely narrow.
signment was not supported by most subsequent authors Scelidosaurus harrisonii Owen, 1861a is known from
(Galton 1973a; Charig & Crompton 1974; Hopson 1975, several partial skeletons from the Lower Lias of Dorset, Eng-
1980). Hopson (1975) demonstrated that Lycorhinus could be land (Early Jurassic: late Sinemurian). Owen (1861a) de-
distinguished from other heterodontosaurids (Abrictosaurus scribed material which he referred to Scelidosaurus, includ-
and Heterodontosaurus). Finally, Gow (1990) referred the ing a femur (GSM 109560), articulated knee-joint (BMNH
maxillae BP/1/4244 (holotype of Lanasaurus Gow, 1975) 39496), ungual phalanx (GSM 10956), a partial juvenile
and BP/1/5253 to Lycorhinus. skeleton (Philpott Museum, Lyme Regis, unnumbered, casts
The validity of Lycorhinus requires reassessment and are catalogued as BMNH R5909) and a near-complete skull
only a preliminary diagnosis is suggested here, based upon (BMNH R1111). Owen (1863) described the near-complete
the marked medial curvature of the dentary and maxillary postcranial skeleton associated with the skull BMNH R1111,
tooth rows (Gow 1990). As discussed by Hopson (1975), a while Lydekker (1888) later designated the articulated knee-
unique combination of plesiomorphic and derived characters joint as the lectotype. Newman (1968) recognised that the
is probably also diagnostic for Lycorhinus. material described by Owen (1861a) represents a composite
Micropachycephalosaurus hongtuyanensis Dong, 1978 of theropod (GSM 109560; BMNH 39496; GSM 10956) and
is known from a partial skull and skeleton (IVPP V5542, ornithischian (Philpott Museum, juvenile skeleton; BMNH
holotype) from the Wangshi Formation of Shandong R1111) material, and Charig & Newman (1992) formally
Province, People’s Republic of China (Late Cretaceous: designated the skull and postcranial skeleton (BMNH R1111)
Campanian). The holotype of Micropachycephalosaurus is as a replacement lectotype. The juvenile described by Owen
(1861a) probably represents a second individual of Scelido- synonymy is based on general similarity, rather than on auta-
saurus (Galton 1975). Further material has come to light in pomorphic features (see Galton 1995); future revision of the
recent years, including BMNH R6704 (Rixon 1968; Charig Thescelosaurus material may indicate the presence of two,
1972: fig. 6A; considered as a possible new taxon of basal or more, distinct taxa. One possible autapomorphy is recog-
thyreophoran by Coombs et al. 1990), BRSMG Ce12785 nised here: a large notch or foramen within the supraoccipital,
(Barrett 2001) and CAMSM X 39256. Reports of the genus dorsal to the foramen magnum (see Galton 1997).
Scelidosaurus in the Kayenta Formation of Arizona (Padian Wannanosaurus yansiensis Hou, 1977 is known from
1989) and the Lower Lufeng of China (Lucas 1996: see Tat- a partial skull and postcranial skeleton (IVPP V4447, holo-
isaurus, below) cannot be substantiated at present. No dia- type) and some referred postcranial elements (IVPP V4447.1,
gnosis based upon synapomorphies has ever been published paratype), from the Xiaoyan Formation of Anhui Province,
for Scelidosaurus and a full diagnosis must await redescrip- China (Late Cretaceous: Campanian). Wannanosaurus is dis-
tion of this taxon (D. B. Norman, unpublished results). tinguished by the extreme flexure of the humerus, with prox-
Scutellosaurus lawleri Colbert, 1981 is known from imal and distal ends set at approximately 30◦ to one another
several partial skeletons (MNA P1.175, holotype; MNA (modified from Sereno 2000).
P1.1752, paratype; for referred material see Rosenbaum & Yandusaurus hongheensis He, 1979 is known from a
Padian 2000) described by Colbert (1981) and Rosenbaum & partial skull and postcranial material (GCC V20501, holo-
Padian (2000) from the Kayenta Formation of Arizona, USA type) from the Upper Shaximiao Formation of Sichuan,
(Early Jurassic: Sinemurian–Pliensbachian). Unique features People’s Republic of China (Late Jurassic: ?Oxfordian,
include: dorsal and ventral margins of the preacetabular pro- Weishampel et al. 2004b). All of the anatomical features pre-
cess of the ilium are drawn out medially into distinct flanges viously used to diagnosis Yandusaurus (He & Cai 1984) have
which converge upon one another anteriorly; elongate tail of wider distributions amongst basal ornithopods. However, one
at least 58 caudal vertebrae (R. J. B. & S. C. R. Maidment, autapomorphic feature is apparent in the holotype (R. J. B.,
unpublished results). pers. obs. 2004; Barrett et al. 2005); the mid–posterior cer-
Stenopelix valdensis Meyer, 1857 is known from a vicals have circular, pit-like depressions developed at the
single partially articulated postcranial skeleton, preserved base of their postzygapophyses.
as impressions in sandstone blocks, from the Obernkirchen Zephyrosaurus schaffi Sues, 1980 is known from an in-
Sandstein of Germany (Early Cretaceous: Berriasian); latex complete skull and postcranial fragments (MCZ 4392, holo-
casts prepared by Sues & Galton (1982) make detailed exam- type) from the Cloverly Formation of Montana, USA (Early
ination of the material possible. The ischium of Stenopelix Cretaceous: Aptian–Albian) and is characterised by the fol-
has the following autapomorphies: distinct bend at mid-shaft; lowing unique combination of characters: prominent boss
broadest at mid-shaft and tapers anteriorly and posteriorly; on anterolateral maxilla; short, massive, triangular palpeb-
blade is transversely arched distally, being ventrally convex ral; boss on jugal; postorbital with distinct projection into or-
and dorsally concave (modified from Sereno 1987). bit; dentary/maxillary teeth with numerous subparallel ridges
Talenkauen santacrucensis Novas et al. 2004 is based connecting to marginal denticles.
upon a partial skull and postcranial skeleton (MPM-10001,
holotype) from the Pari Aike Formation, Santa Cruz Excluded species level taxa
Province, Argentina (Late Cretaceous: Maastrichtian). This A number of taxa have been considered non-diagnosable
taxon can be diagnosed by the presence of well-developed nomina dubia by most recent reviews and are here excluded
epipophyses on cervical 3 and plate-like uncinate processes from analysis. These taxa include ‘Camptosaurus’ leedsi,
on the rib-cage (Novas et al. 2004). Fabrosaurus australis, Geranosaurus atavus, ‘Hypsilopho-
The genus Tenontosaurus Ostrom, 1970 is character- don’ wielandi, Laosaurus celer, Laosaurus minimus, Lus-
ised by the following features: dorsoventrally tall maxilla, itanosaurus liasicus, Nanosaurus agilis and Sanpasaurus
nearly full height of the rostrum; orbit square; 12 cervical yaoi.
vertebrae; elongate tail (59+ caudals). Two species are re- Many ornithischian or putative ornithischian taxa have
cognised and included here: Tenontosaurus tilletti Ostrom, been erected solely or largely on the basis of dental re-
1970 is known from abundant material (see Forster 1990) mains, including: Alocodon kuehnei, Crosbysaurus harri-
from the Cloverly Formation of Montana, USA (Early Creta- sae, Drinker nisti, Ferganocephale adenticulatum, Galto-
ceous: Aptian–Albian) and Tenontosaurus dossi Winkler nia gibbidens, Gongbusaurus shiyii, Krzyzanowskisaurus
et al. 1997 is known from two specimens from the Twin hunti, Lucianosaurus wildi, Pekinosaurus olseni, Phyllodon
Mountains Formation of Texas, USA (Early Cretaceous). henkeli, Protecovasaurus lucasi, Revueltosaurus callenderi,
The species are distinguished by the retention in Tenonto- Siluosaurus zhangqiani, ‘Stegosaurus’ madagascariensis,
saurus dossi of premaxillary teeth and a postpubic process Taveirosaurus costai, Tecovasaurus murryi and Trimucrodon
equal in length to the ischium (Winkler et al. 1997). cuneatus. Recent work has demonstrated that at least some
Thescelosaurus neglectus Gilmore, 1913 is known of these taxa pertain to non-ornithischian clades (Parker et al.
from numerous specimens (see Galton 1997) from the 2005a; Irmis et al. 2007) and the taxonomic validity of many
Lance Formation of Wyoming, the Hell Creek Formation of of these tooth taxa is additionally questionable (Weishampel
Montana, the Scollard Formation of Alberta and the French- et al. 2004a). Furthermore, tooth taxa add little new anatom-
man Formation of Saskatchewan (Late Cretaceous: ?late ical information to the analysis and suffer from extremely
Campanian–Maastrichtian). The taxonomic history of Thes- high (more than 95%) levels of missing data. For these reas-
celosaurus has been summarised by Galton (1995, 1997) ons these taxa have been excluded.
and his taxonomic assignments are followed provisionally Technosaurus smalli, from the Cooper Canyon Form-
here, with Thescelosaurus edmontonensis being considered ation (Late Triassic: Norian) of Texas, was described as a
a junior synonym of T. neglectus. It should be noted that ‘fabrosaurid’ ornithischian by Chatterjee (1984) on the basis
of a single fragmentary skeleton. Sereno (1991a) sugges- taxa may prove to be diagnostic with further study, we here
ted that the holotype of Technosaurus contains elements of provisionally exclude them from phylogenetic analysis as
both an ornithischian and a sauropomorph. Irmis et al. (2005, we have been unable to examine the majority of the material
2007) agreed that the specimen is a composite of at least two first-hand.
taxa, but suggested that the posterior portion of the lower jaw A number of taxa (Auroraceratops rugosus,
is referable to the pseudosuchian Shuvosaurus, whereas other Changchunsaurus parvus, Xuanhuaceratops niei, Yamacer-
parts of the holotype may represent a Silesaurus-like taxon. atops dorngobiensis, Yinlong downsi) were described after
They do not consider any of the material to be ornithischian. the current analysis was carried out and thus have not been
In view of the considerable confusion as to the association included. We plan to include these taxa in future iterations
of the holotype of Technosaurus, it is here excluded from of this analysis.
phylogenetic analysis.
Norman et al. (2004b) considered Tatisaurus oehleri
and Bienosaurus lufengensis from the Lower Lufeng Form-
Selection of outgroup taxa
ation (Early Jurassic: Sinemurian) of China as valid taxa Three outgroup taxa were chosen, based upon the phylo-
and possible basal thyreophorans. However, reassessment of genetic framework outlined above. Recent, comprehens-
Tatisaurus (Norman et al. 2007) suggests that it is a no- ive, phylogenetic analyses of basal dinosaurs have demon-
men dubium. Autapomorphic characters are not evident in strated that Herrerasaurus ischigualastensis Reig, 1963 is
the original description of Bienosaurus (Dong 2001) and the the most basal known member of Saurischia (Langer 2004;
validity of this taxon is uncertain. As a result, both taxa are Langer & Benton 2006). In addition, this taxon is known
excluded from the current phylogenetic analysis. from well-preserved complete material and has been extens-
Xiaosaurus dashanpensis from the Lower Shaximiao ively described (Novas 1993; Sereno 1993; Sereno & Novas
Formation of Sichuan Province, China (Middle Jurassic: 1993). Marasuchus talampayensis (Romer, 1972) represents
?Bajocian, Chen et al. 1982; ?Bathonian–Callovian, Dong a well-known dinosaurian outgroup and Euparkeria capensis
& Tang 1984) is based upon very fragmentary remains, but Broom, 1913 represents a basal archosaur, phylogenetically
appears to be diagnosable on the basis of the possession of a distant from Dinosauria and lacking the numerous derived
proximally straight humerus that lacks the medial curvature specialisations seen in many other more proximate dinosaur-
seen in all other basal ornithischians (Barrett et al. 2005). ian outgroups, such as pterosaurs and crurotarsans.
However, the whereabouts of the holotype and referred ma-
terial are currently unknown (Barrett et al. 2005) and un-
available for further study and the original description (Dong
& Tang 1983) provides few anatomical details. As a result,
Analyses
we do not include Xiaosaurus in the present study.
The location of the holotype of ‘Gongbusaurus’ wu-
Search methods
caiwanensis is currently unknown (X. Xu, pers. comm., The full matrix (Appendix 3) consists of 46 taxa (43 in-
2004) and the original description (Dong 1989) is brief and group taxa and 3 outgroup taxa), coded for 221 characters
poorly figured and does not allow the recognition of auta- (Appendix 2). The data matrix was constructed using the
pomorphies, or a unique character combination. Therefore, NEXUS Data Editor (http://taxonomy.zoology.gla.ac.uk/rod/
despite the fact that recent reviews (e.g. Norman et al. 2004c) NDE/nde.html). Prior to analysis, safe taxonomic reduc-
retain ‘Gongbusaurus’ wucaiwanensis as a valid taxon, we tion (Wilkinson 1995c) was carried out using the pro-
here exclude it from the phylogenetic analysis. gram TAXEQ3 (Wilkinson 2001a). Safe taxonomic reduc-
Notohypsilophodon comodorensis was described by tion identifies taxa that can be excluded without affecting
Martı́nez (1998) on the basis of a partial skeleton from the the inferred relationships of the remaining taxa. The matrix
Bajo Barreal Formation, Chubut Province, Argentina (Late does not contain any taxonomic equivalents and all taxa were
Cretaceous: ?Cenomanian). Martı́nez (1998) listed a number included in subsequent analyses.
of potential autapomorphies of Notohysilophodon; however, Analyses were carried out in PAUP∗ 4.0b10 (Swofford
most of these appear to have wider distributions within Or- 2002); all characters are treated as unordered and equally
nithischia (e.g. the reduction of the deltopectoral crest of the weighted and polymorphisms are treated as uncertainty.
humerus is seen in other South American ornithopods, Novas Branches with a minimum length of zero were collapsed dur-
et al. 2004) or represent plesiomorphies (distal end of fibula ing searches (the ‘-amb’ option); this setting recovers only
reduced, astragalus with ‘stepped’ proximal surface). Noto- ‘strictly supported’ trees (Nixon & Carpenter 1996; Kearney
hypsilophodon may represent a valid taxon, but this cannot & Clark 2003), but can result in trees that are not of min-
be ascertained from the published description and this taxon imum length and cannot be considered as most parsimonious
is provisionally excluded from the phylogenetic analysis. trees (MPTs); (Wilkinson 1995a). As a result we filtered
Four taxa (Atlascopcosaurus loadsi, Fulgurotherium the resultant set of trees to ensure that only minimum length
australe, Leaellynasaura amicagraphica, Quantassaurus in- trees were retained. Analysis was conducted using a heuristic
trepidus) have been named on the basis of cranial and post- search with 10,000 replicates and TBR branch-swapping,
cranial material from the Early Cretaceous of Australia (Rich each starting tree being produced by random stepwise addi-
& Vickers-Rich 1989, 1999). Although all of these taxa have tion.
been considered valid by a recent review (e.g. Norman et al. The analysis recovered 3787 trees; filtering these trees
2004c), the type specimens of all four taxa are fragmentary so that only minimum length trees were retained resulted
and unambiguous autapomorphies have not yet been defined. in 756 MPTs of 477 steps (Consistency Index (CI) = 0.505,
Referral of additional material to any one of these taxa is Retention Index (RI) = 0.732, Rescaled Consistency Index
problematic. Although we accept that some or all of these (RC) = 0.370). Strict and 50% majority-rule component
Figure 2 Strict component consensus (left) and 50% majority-rule consensus (right) of 756 most parsimonious trees (MPTs) produced by
analysing a data matrix of 46 taxa and 221 characters. Values above nodes on the Strict component consensus represent bootstrap proportions.
Values beneath nodes on the Strict component consensus indicate Bremer support. Bremer support values of +1 or less are not shown. Numbers
beneath nodes on the 50% majority-rule consensus indicate the percentage of MPTs in which that node appears (nodes with no values beneath
them appear in all MPTs).
consensus trees (Fig. 2) and an Adams consensus tree, were amounts of missing data, or character conflict, or both. A
calculated using PAUP∗ . The strict component consensus maximum agreement subtree was also calculated that ex-
(SCC) tree contains two major polytomies and contains a cludes 8 taxa (Echinodon, Lycorhinus, Bugenasaura, Jeholo-
much lower degree of resolution than the majority-rule or saurus, Talekauen, Thescelosaurus, Yandusaurus, Zephyro-
Adams consensus trees; the latter observation suggests that saurus; see Fig. 3).
the low degree of resolution in the SCC tree results from An additional search was carried out using a demonstra-
a number of taxa acting as ‘wildcards’, as a result of high tion version of TNT (Tree Analysis Using New Technology)
Figure 3 Maximum agreement subtree of 756 most parsimonious trees (MPTs) produced by analysing a data matrix of 46 taxa and 221
characters. Eight (Echinodon, Lycorhinus, Bugenasaura, Jeholosaurus, Talekauen, Thescelosaurus, Yandusaurus and Zephyrosaurus) of the
original taxa are excluded.
v1.0, downloaded from www.zmuc.dk/public/phylogeny. A The set of 756 MPTs recovered by PAUP∗ forms the basis
‘New Technology’ search was carried out, using a random for subsequent discussion.
addition-sequence, 1000 replicates and default settings for In an attempt to resolve further relationships com-
the ‘Sect. Search’, ‘Ratchet’, ‘Drift’ and ‘Tree Fusing’ op- mon to all 756 MPTs and to identify the most unstable
tions. The search recovered 119 trees of 477 steps; the con- taxa, reduced consensus techniques (Wilkinson 1994, 1995b,
sensus of these trees matched the consensus of the 756 MPTs 2003) were applied to the data. The most commonly used
recovered by PAUP∗ . That TNT failed to find trees shorter consensus methods are strict component consensus (SCC)
than 477 steps suggests that this is the minimum tree length. trees, which include all terminal taxa and all the clades
(components) common to all MPTs. However, the strict com- found only within the clades Thyreophora and Iguanodontia,
ponent method has problems of insensitivity and may fail to but even here support is relatively low. However, it is possible
represent relationships that are common to the set of MPTs, that a few unstable ‘wildcard’ taxa, such as those identified
but cannot be expressed as shared clades (Wilkinson 2003). by reduced consensus techniques (see above), can obscure
Reduced consensus methods identify ‘n-taxon statements’; support for relationships, resulting in lower decay indices
n-taxon statements express cladistic relationships (e.g. A and than might be otherwise expected.
C are more closely related to each other than either is to E), Wilkinson et al. (2000) proposed a new technique,
but need not include all terminal taxa. Reduced consensus double decay analysis (DDA), which provides Bremer sup-
methods delete unstable taxa to produce more informative port for all strictly supported n-taxa relationships. An attempt
consensus trees, which represent n-taxon statements. was made to apply DDA using RADCON (Thorley & Page
Reduced consensus was applied using the ‘strict’ pro- 2000); however, the large size of the data set meant that
gram of REDCON 3.0 (Wilkinson, 2001b) and the results this approach was not feasible due to time and memory con-
corroborated using RADCON (Thorley & Page 2000), identi- straints. For particular areas of interest PAUP∗ was used to
fying a profile of eight strict reduced consensus (SRC) trees, write backbone constraints that could then be used to test
the first of which includes all taxa and corresponds in to- the decay indices of n-taxa statements (see individual ex-
pology to the SCC tree of the 756 MPTs. The remaining amples below). This allowed an assessment of the effect that
SRC trees exclude one or more unstable ‘wildcard’ taxa, res- wildcard taxa have upon the Bremer support for clades.
ulting in an increase in resolution. Seven taxa (Echinodon,
Lycorhinus, Zephyrosaurus, Talenkauen, Yandusaurus, Gas-
parinisaura, Parksosaurus) are identified as unstable by these Bootstrapping
analyses. We combined six of the SRC trees (those exclud- Bootstrap analysis generates ‘pseudoreplicate’ data sets by
ing Echinodon, Lycorhinus, Zephyrosaurus, Talenkauen and randomly sampling with replacement a proportion of the
Yandusaurus) to produce an informative derivative SRC tree characters, deleting some characters randomly and reweight-
(Fig. 4). We use this derivative SRC tree as the basis for ing other characters randomly. The MPTs are generated for
optimisation of synapomorphies (Appendix 4) and for much each pseudoreplicate and the degree of conflict between res-
of the subsequent discussion, and it represents our preferred ulting MPTs is assessed using a 50% majority rule consensus
hypothesis of interrelationships. tree. Clades that are supported by a large number of charac-
ters, with low levels of homoplasy, would be expected to have
high bootstrap values, whereas bootstrap values should be
Testing the support for relationships lower for clades supported by only a few, or by homoplastic,
Randomisation tests characters. The bootstrap should, therefore, be considered as
PAUP∗ was used to run a Permutation Tail-Probability (PTP) a one-sided test of a cladogram (Kitching et al. 1998): groups
test using 1000 randomised replicates of the reduced data that are recovered are supported by the data, but groups that
set (Faith & Cranston 1991; Kitching et al. 1998). The ran- are not recovered (or that have low bootstrap values) cannot
domised replicates are created by randomly permuting the be rejected.
character states assigned to taxa, decreasing character con- A bootstrap analysis was carried out using PAUP∗ with
gruence to a level that would be expected by chance alone. 1000 replications. To allow the analysis to be carried out
The MPT length is then calculated for each replicate and the within a reasonable length of time the MAXTREES option
distribution of MPT lengths for the replicates is compared of PAUP was set to 1000. This means that for each pseu-
to the length of the original MPT. The PTP test has been doreplicate data set the search for MPTs was truncated once
criticised (Bryant 1992; Carpenter 1992) and Kitching et al. 1000 trees had been found. Figure 2 shows the results of
(1998) suggested that it could best be used as an independent the bootstrap analysis. Bootstrap support is weak throughout
evaluation of the explanatory power of the data set, rather much of the tree.
than as a criterion for acceptance or rejection of any partic- Unstable ‘wildcard’ taxa can obscure levels of bootstrap
ular cladogram. In this case, the results of this test indicate support for relationships (Wilkinson 2003). One potential
that the most parsimonious tree length (477 steps) lies out- solution is to use the majority-rule bootstrap reduced con-
side the range of minimum tree lengths obtained from the sensus (MBRC) technique developed by Wilkinson (1996).
randomised data (P = 0.001). This indicates that a signific- The MBRC method provides bootstrap proportions for all
ant phylogenetic signal is present in the data set and is not ‘n-taxon statements’ and is implemented in the REDBOOT
completely obscured by character conflict and missing data. program of REDCON 3.0. Unfortunately, REDBOOT could
not be used for this analysis due to the large size of the
data matrix. An alternative method for assessing the im-
Bremer support pact of ‘wildcards’ on bootstrap support is used here. All
‘Traditional’ decay analysis, or Bremer support, measures trees recovered by the bootstrap analysis were saved to a
the number of additional steps required before the clade is NEXUS treefile. The five unstable taxa removed in the de-
lost from the strict consensus of near-minimum length clado- rivative SRC tree (Echinodon, Lycorhinus, Zephyrosaurus,
grams (Bremer 1988; Kitching et al. 1998). Bremer support Talenkauen and Yandusaurus) were pruned a posteriori from
was calculated for nodes present in the SCC tree by search- this set of bootstrap trees, with duplicate topologies cre-
ing in PAUP∗ for the shortest trees not compatible with a ated by this deletion being collapsed. Following pruning of
particular node, using the CONVERSE option. these five taxa a new majority-rule bootstrap reduced con-
Bremer support values are shown in Fig. 2. Most nodes sensus tree was generated containing recalculated bootstrap
have a decay index of +1, i.e. they are absent from the strict values (Fig. 4). For further areas of particular interest addi-
consensus of all trees of 478 steps or less. Stronger support is tional potentially unstable taxa were identified and pruned
Figure 4 Derivative strict reduced consensus tree derived by a posteriori pruning of five unstable taxa (Echinodon, Lycorhinus,
Zephyrosaurus, Talenkauen and Yandusaurus) from the set of 756 most parsimonious trees (MPTs) generated by the full analysis. The number
above each node is a unique identifier used in the tree description (see Appendix 4). The number beneath a node represents the bootstrap
proportion for that node (taken from the reduced bootstrap analysis). Note the increased levels of bootstrap support for a number of nodes
when compared to Fig. 2. Abbreviations: ORN., Ornithischia; HETERODONT., Heterodontosauridae; GENA., Genasauria; NEORN., Neornithischia;
ORNITH., Ornithopoda; MARG., Marginocephalia; CERAT., Ceratopsia; PACHY., Pachycephalosauria.
and bootstrap values recalculated. The effect of ‘wildcards’ Some previously suggested ornithischian synapo-
on bootstrap values is discussed further below. morphies are not supported by this analysis. For example
Sereno (1999a) suggested that an elongate posterolateral
process of the premaxilla diagnoses the clade; however,
an elongate posterolateral process is present in the basal
Results archosaur Euparkeria (Ewer 1965), the basal saurischian
Herrerasaurus (Sereno & Novas 1993) and the problem-
Ornithischian monophyly atic non-ornithischian ‘dinosauriform’ Silesaurus opolen-
Most of the characters identified as synapomorphic for Or- sis (Dzik 2003), and may be plesiomorphic for Dinosauria
nithischia (Appendix 4) have been identified by previous (Maryańska & Osmólska 1985).
authors (Norman 1984; Sereno 1984, 1986, 1991a, 1999a; Many of the characters identified by this and previ-
Cooper 1985; Maryańska & Osmólska 1985; Weishampel ous analyses as synapomorphic for Ornithischia describe the
2004; Norman et al. 2004a; Butler 2005). Some new potential anatomy of the dentary and maxillary teeth. These synapo-
ornithischian synapomorphies are proposed by this analysis. morphies have been used to refer taxa named on the basis
The presence of a buccal emargination has previously been of isolated teeth to Ornithischia (e.g. Hunt & Lucas 1994).
considered (Sereno 1986, 1999a) to be synapomorphic for a However, recent discoveries suggest that ornithischian-like
less inclusive clade of ornithischians, Genasauria, and absent teeth have evolved a number of times within Archosauria.
in the basal ornithischian Lesothosaurus diagnosticus. This For instance, recently discovered cranial and postcranial ma-
analysis alternatively suggests that the presence of a weak or terial of the putative ornithischian Revueltosaurus callenderi
incipient buccal emargination (generally correlated with the suggests that this taxon is actually a non-ornithischian archo-
presence of ‘cheeks’, see Galton 1973a) is a synapomorphy saur that appears to be phylogenetically closer to crocodylo-
of Ornithischia. As noted by Butler (2005: 204), in Lesotho- morphs than to dinosaurs and that its ornithischian-like den-
saurus there is a weak anteroposteriorly extending ridge (re- tition evolved independently (Parker et al. 2005a, b; Irmis
ferred to, below, as the ‘maxillary ridge’) above the row of ex- et al. 2007). In addition, ornithischian-like teeth have been
ternal maxillary foramina, forming the ventral margin of the described in Silesaurus opolensis (Dzik 2003), while many
external antorbital fenestra. Below this eminence the lateral of the characteristic features of the ornithischian dentition
surface of the maxilla is gently bevelled such that the maxil- also occur in taxa as diverse as therizinosaurs, basal sauro-
lary tooth row is slightly inset along at least the posterior two- podomorphs and aetosaurs, suggesting that dental characters
thirds of its length. A similar weak medial offset of the tooth (perhaps not surprisingly) may be subject to particularly high
row is seen in other basal ornithischians such as Abricto- levels of homoplasy.
saurus consors (BMNH RUB54) and Scutellosaurus lawl- The identification of ornithischian-like dental character
eri (Colbert 1981) and is proposed here to be homologous states in non-ornithischian taxa highlights the problems in-
with the well-developed buccal emargination seen in many herent in referring fragmentary material to specific clades.
other ornithischians. Irmis et al. (2007) criticised Butler As noted by Butler et al. (2006) and Irmis et al. (2007), most
(2005) for using the same coding for the weak buccal Triassic taxa named upon the basis of isolated teeth cannot,
emargination of Lesothosaurus and the well-defined buccal in general, be referred to Ornithischia with certainty unless
emargination of taxa such as Heterodontosaurus tucki. How- they demonstrate, or are associated with material demon-
ever, the coding of Butler (2005) referred only to the presence strating, unique synapomorphies of an ornithischian clade
of an emargination (not how well-developed it was), which (i.e. features that are not independently synapomorphic for
we propose is homologous in Lesothosaurus and Hetero- other clades).
dontosaurus. Irmis et al. (2007) additionally stated that in
ornithischians the maxillary ridge was separated from the
ventral margin of the external antorbital fenestra. This is true
Pisanosaurus mertii
for some ornithischians (e.g. Hypsilophodon foxii, Galton Pisanosaurus mertii is generally believed to be the oldest
1974a) but in many basal ornithischians such as Lesotho- known ornithischian (Casamiquela 1967), but both the as-
saurus (BMNH R8501) and Heterodontosaurus (SAM-PK- sociation of the material included within the holotype speci-
K1332; Norman et al. 2004c: fig. 18.1) the maxillary ridge men and its phylogenetic position, have proved controversial.
does form the ventral margin of the external antorbital fen- Sereno (1991a) proposed that the holotype specimen con-
estra. tained material from more than one individual; he suggested
Another potential ornithischian synapomorphy is the that the skull fragments, partial impression of the pelvis and
size and position of the posttemporal foramen. In ornith- distal hind limb might belong together, but that the fragment-
ischian outgroups the posttemporal foramen is relatively ary scapula and other postcranial bones were too small to be
large and positioned on the boundary between the parietal referred to the same individual. However, Bonaparte (1976)
and the paroccipital process (e.g. Euparkeria capensis, Ewer noted that the vertebrae were recovered in articulation with
1965: fig. 2B; basal sauropodomorphs, Galton & Upchurch the skull and that the skeleton may have been complete prior
2004a; Herrerasaurus ischigualastensis, Sereno & Novas to weathering, and a recent review supported the idea that
1993: fig. 8C), whereas in basal ornithischians (e.g. Lesotho- the type specimen represents a single individual (Irmis et al.
saurus diagnosticus, Sereno 1991a, Sereno 1991a: fig. 11C; 2007).
Heterodontosaurus tucki, Weishampel & Witmer 1990b: fig. Bonaparte (1976) referred Pisanosaurus to Heterodon-
23.1), the posttemporal foramen is reduced in size and en- tosauridae on the basis that both share subcylindrical, closely
tirely enclosed by the paroccipital process. This character was packed cheek teeth, with wear facets forming a more or
independently identified as an ornithischian synapomorphy less continuous surface extending along the tooth row. How-
by Langer & Benton (2006). ever, Weishampel & Witmer (1990a) and Sereno (1991a)
considered Pisanosaurus to be the most basal known or- the result of the unstable and problematic taxon Echinodon
nithischian. Weishampel & Witmer (1990a) suggested that (which is highly fragmentary, with 83% missing data); ex-
the similarities between Pisanosaurus and heterodontosaur- clusion of Echinodon results in the remaining four OTUs
ids are plesiomorphic, while Sereno (1991a: 174) noted that: forming a heterodontosaurid clade, while further exclusion
‘The [wear] facets . . . do not form a continuous occlusal sur- of Lycorhinus (86% missing data) in the derivative SRC tree
face as occurs in Heterodontosaurus [tucki].’ More recently, resolves relationships within Heterodontosauridae (Fig. 4).
Norman et al. (2004a) have once again emphasised the mor- The heterodontosaurid node is weakly supported by
phological similarities between the cranial material of Pis- bootstrap proportions in the total-evidence bootstrap ana-
anosaurus and that of heterodontosaurids. lysis (support of only 19%); however, bootstrap support is
The SCC tree (Fig. 2) recovered by this analysis posi- considerably higher (68%) in the reduced bootstrap analysis
tions Pisanosaurus in an unresolved polytomy at the base of (Fig. 4), suggesting that low bootstrap support for Heterodon-
Ornithischia. However, the 50% majority-rule consensus tree tosauridae is probably the result of fragmentary and unstable
(Fig. 2), maximum agreement subtree (Fig. 3) and the derivat- wildcard taxa. To test whether similar ‘hidden’ Bremer sup-
ive SRC tree (Fig. 4) all support the position of Pisanosaurus port exists for Heterodontosauridae we wrote a backbone
as the most basal known ornithischian. These consensus trees constraint that specified that Heterodontosaurus, Abricto-
additionally position heterodontosaurids as a monophyletic saurus and BMNH A100 form a clade to the exclusion of
clade of non-genasaurians, close to the base of Ornithischia other ornithischians, but did not specify the position of the
(discussed below). Pisanosaurus and heterodontosaurids are five unstable taxa (Echinodon, Lycorhinus, Zephyrosaurus,
not separated in the SCC (this is the result of the instability of Talenkauen and Yandusaurus) discussed above and removed
the fragmentary wildcard taxon Echinodon becklesii); only in the derivative SRC tree. The Bremer support for this back-
one node separates Pisanosaurus from heterodontosaurids in bone constraint was +2, again suggesting that low support
the derivative SRC tree. This node is weakly supported by for Heterodontosauridae is the result of unstable wildcard
bootstrap proportions: it does not appear in the total-evidence taxa.
bootstrap analysis (support of only 37%) and has support of The MPTs recovered in this analysis do not support
only 54% in the reduced bootstrap analysis (Fig. 4). Only one a link between heterodontosaurids and ornithopods (phylo-
character (Character 206, Appendix 2) unambiguously sup- genetic hypothesis 1, above). In order to test this further we
ports this node. Constraining Pisanosaurus and heterodon- ran the following constrained analyses: firstly, Ornithopoda
tosaurids to form a monophyletic clade requires only one ad- (sensu Weishampel 1990; Sereno 1999a), containing hetero-
ditional step. This suggests that the two opposing phylogen- dontosaurids, ‘hypsilophodontids’ and iguanodontids, was
etic positions for Pisanosaurus, as either a non-genasaurian constrained as monophyletic, although Hypsilophodontidae
basal ornithischian (Sereno 1991a) or a heterodontosaurid (sensu Weishampel & Heinrich 1992) was not constrained as
(Bonaparte 1976), are not necessarily mutually exclusive. a monophyletic clade; secondly, a backbone constraint was
Pisanosaurus may indeed represent a heterodontosaurid and specified that required that Heterodontosaurus and Ankylo-
the evidence for this should be reconsidered by future work. pollexia be more closely related to each other than either
is to marginocephalians, thyreophorans or ornithischian out-
groups. This constraint does not specify the complete content
Heterodontosauridae of Ornithopoda.
The position of heterodontosaurids within Ornithischia is one Templeton non-parametric tests were carried out using
of the most problematic areas in ornithischian phylogeny and PAUP∗ that compared trees within the profile of 756 MPTs
there is no current consensus on this topic. Four alternative generated by the unconstrained analysis with all trees re-
phylogenetic positions have been proposed: (1) as basal or- covered by the constrained analyses. Ideally all MPTs should
nithopods (e.g. Crompton & Charig 1962; Thulborn 1971; be compared with all trees recovered by the constrained ana-
Galton 1972; Santa Luca et al. 1976; Sereno 1984, 1986, lyses; however, as this is time intensive (this would involve
1999a; Gauthier 1986; Weishampel 1990; Weishampel & 756 separate Templeton tests) a subset of the MPTs (every
Witmer 1990b; Smith 1997; Norman et al. 2004c); (2) as 25th MPT) was used.
the sister group to Marginocephalia (Maryańska & Osmólska Under the first constraint, 1812 trees of 500 steps were
1984; Cooper 1985; Olshevsky 1991; Zhao et al. 1999; Buch- found, 23 steps longer than the most parsimonious tree
holz 2002; You et al. 2003; Norman et al. 2004c; Xu et al. length. These trees were a significantly worse (p = 0.0004–
2006); (3) as the sister group to Ornithopoda + Margino- 0.01) explanation of the data than the most parsimonious
cephalia (Cerapoda) (Norman 1984; Maryańska & Osmólska topology. The second constraint recovered 2034 trees of 490
1985; Butler 2005); (4) as the most basal well-known ornith- steps (13 extra steps). Some, but not all, of these trees were
ischians (Bakker & Galton 1974; Olsen & Baird 1986). This significantly (p = 0.03–0.07) worse explanations of the data
analysis supports the fourth of these positions and the phylo- than the most parsimonious topology.
genetic support for this is discussed below, although a full A sister group relationship between heterodontosaurids
review of the anatomical evidence will be presented else- and marginocephalians (phylogenetic hypothesis 2, above)
where (R. J. B., unpublished results). is not found by this analysis; instead heterodontosaurids are
Four taxa (Heterodontosaurus tucki, Abrictosaurus con- positioned more basally within Ornithischia (Fig. 4). To test if
sors, Echinodon becklesii, Lycorhinus angustidens) and one any support is present in the data for a Heterodontosauridae–
specimen (BMNH A100) previously referred to Heterodon- Marginocephalia sister grouping we constrained such a clade
tosauridae were included in this analysis. These taxa do not to be monophyletic and 420 trees of 487 steps (10 extra steps)
form a clade in the SCC tree, but are included in a large were recovered; Templeton tests indicate that these trees are
polytomy at the base of Ornithischia (Fig. 2). However, re- not a significantly worse explanation of the data (p = 0.128–
duced consensus trees indicate that this basal polytomy is 0.1803).
A number of derived character states are absent in het- basal neornithischians. Of these characters, reduction of the
erodontosaurids, but occur in all cerapodan ornithischians. forelimb (character 153, Appendix 2) is here optimised as
These include: loss of the squamosal–quadratojugal con- independently gained in Lesothosaurus, Agilisaurus louder-
tact; closure of the external mandibular fenestra (reversed backi and pachycephalosaurs; the presence of a dorsal groove
in some psittacosaurids); reduction of the pubic peduncle on the ischium (character 183, Appendix 2) may be a gena-
of the ilium; development of an elongated rod-like prepubic saurian plesiomorphy as it appears to be present in the thyreo-
process; modification of the fossa trochanteris into a dis- phoran Scutellosaurus lawleri (UCMP 130580, R. J. B., pers.
tinct constriction separating the femoral head and greater obs., 2005); while the reduction of metatarsal one (char-
trochanter; and the anteroposterior expansion of the greater acter 211, Appendix 2) optimises as an ornithischian ple-
trochanter of the femur relative to the anterior trochanter siomorphy, present in heterodontosaurids.
(Characters 52, 104, 178, 193, 194, 198, 199: Appendix 2). If This analysis resolves Lesothosaurus as the most basal
heterodontosaurids are positioned within Cerapoda, as either thyreophoran (Figs 2–4; see also Liu 2004); however, only
basal marginocephalians or basal ornithopods, then a sub- one unambiguous character (character 106, Appendix 2) sup-
stantial amount of homoplasy must be invoked to explain the ports this position. In Lesothosaurus an anteroposteriorly ex-
distribution of these derived character states. tending ridge is present on the lateral surface of the surangu-
This analysis finds support for an alternative position, lar, immediately anterodorsal to the glenoid and dorsal to the
not suggested by previous cladistic phylogenetic analyses surangular foramen (BMNH RUB17; a previous reconstruc-
(although suggested by the non-cladistic work of Bakker & tion (Sereno 1991a) shows this feature as less prominent than
Galton 1974; Olsen & Baird 1986), that heterodontosaurids actually occurs in the syntype and referred specimens). This
are not members of Genasauria and are actually situated close feature is also present in the basal thyreophorans Scelido-
to the base of Ornithischia. This basal position is supported by saurus (BMNH R1111), Emausaurus (Haubold 1990) and
a number of characters (Appendix 4), including: absence of Scutellosaurus (UCMP 130580). In all four taxa the ridge is
a spout-shaped mandibular symphysis; premaxillary crowns prominent and transversely thickened, overhangs the lateral
not expanded mesiodistally or apicobasally above the root; al- surface of the angular and laterally delimits a narrow, dorsally
veolar foramina absent medial to maxillary and dentary tooth facing shelf. Statistical support for this position of Leso-
rows; retention of epipophyses on cervical vertebrae; manus thosaurus is weak (total-evidence bootstrap proportion =
length more than 40% of the combined length of the humerus 40%; Bremer support = +1), with the low bootstrap support
and radius; penultimate phalanges of the second and third reflecting the fact that this node is supported by only a single
manual digits more elongate than the proximal phalanges; character.
extensor pits present on the dorsal surface of the distal end of All previous and current interpretations of the exact
metacarpals and manual phalanges; manual unguals strongly phylogenetic position of Lesothosaurus are problematic and
recurved with prominent flexor tubercles (Characters 97, 113, poorly supported. However, it seems clear that Lesothosaurus
126, 133, 156, 159, 162, 163: Appendix 2). is positioned close to the base of Genasauria, as either the
A basal position for heterodontosaurids is more con- sister-taxon to this clade (Sereno 1986, 1999a), the most
sistent with the stratigraphic record than previous hypotheses basal known neornithischian (Butler 2005), or the most basal
and has important implications for our understanding of early thyreophoran (this analysis). In addition, autapomorphies ap-
ornithischian anatomy, palaeobiology and evolution that will pear to be difficult to delimit for Lesothosaurus (Butler 2005).
be explored by future work. However, substantial further Taken in combination, these observations suggest that the
work is required on heterodontosaurid anatomy and char- anatomy of Lesothosaurus may be close to that of the ances-
acter homology before this hypothesis can be considered tral genasaurian and that this taxon remains a good ‘ancestral’
well-supported. taxon for ornithischian functional studies.
A monophyletic Fabrosauridae was not supported by
this analysis. Peng (1997) identified a number of characters
‘Fabrosaurids’ that he suggested linked Lesothosaurus and Agilisaurus to
Sereno (1984, 1986, 1991a, 1999a) proposed that the ple- the exclusion of other ornithischians. While some of these
siomorphic ornithischian Lesothosaurus diagnosticus is the characters have more widespread distributions within Or-
sister group of Genasauria, a position similar to that pro- nithischia (e.g. Characters 173, 176, 183, 211, Appendix 2),
posed by Galton (1972) and this interpretation has been fol- one character (humerus strongly reduced in length, Char-
lowed by later authors (e.g. Weishampel & Witmer 1990a; acter 153, Appendix 2) is only known in Lesothosaurus
Norman et al. 2004a). This contrasts with previous views and Agilisaurus amongst basal ornithischians. This character,
of ornithischian phylogeny (Thulborn 1971; Norman 1984; therefore, provides some evidence for a monophyletic Fab-
Cooper 1985; Maryańska & Osmólska 1985) that usually rosauridae. However, Agilisaurus is more derived than Leso-
considered Lesothosaurus to have affinities with Ornitho- thosaurus in a large number of other character states (e.g.
poda and, sometimes, a larger group that additionally in- Characters 101, 104, 178, 184, 194, Appendix 2), suggesting
cluded pachycephalosaurs and ceratopsians (Norman 1984; that Fabrosauridae sensu Peng (1997) is defined largely on
Butler 2005). The interpretation of the position of Lesotho- the basis of symplesiomorphies and is probably paraphyl-
saurus suggested by Sereno (1984, 1986, 1991a, 1999a) etic. Five additional steps are required for Lesothosaurus
is based upon a number of putative synapomorphies sup- and Agilisaurus to form a clade to the exclusion of other
posedly shared by Genasauria and absent in Lesothosaurus; taxa.
Butler (2005) recently demonstrated that most of these char- Neither Butler (2005) nor this analysis have been able to
acters are either present in Lesothosaurus or absent in some find anatomical evidence for a South African ‘fabrosaurid’
basal thyreophorans/neornithischians. Butler (2005) sugges- clade (sensu Knoll 2002a, b), consisting of Lesothosaurus
ted three characters that might group Lesothosaurus with and Stormbergia dangershoeki, although only one additional
step is required for such a clade to be recovered. One un- not a significantly worse explanation of the data (Templeton
ambiguous synapomorphy positions Stormbergia as a basal test, p = 0.04–0.13).
neornithischian, separate from Lesothosaurus: presence of a
tab-shaped obturator process on the ischium (Character 184,
Appendix 2).
Basal neornithischians
Most previous analyses have included all neornithischian
taxa within Cerapoda, as members of either Ornithopoda or
Thyreophora Marginocephalia (e.g. Sereno 1999a). Only Butler (2005) has
The monophyly of Thyreophora (ignoring, at present, the previously identified non-cerapodan neornithischians. This
possible most basal thyreophoran Lesothosaurus diagnos- analysis finds four OTUs (Stormbergia dangershoeki, Agil-
ticus, discussed above) is well-established and uncontro- isaurus louderbacki, Hexinlusaurus multidens, Othnielia rex;
versial and the clade including Scutellosaurus lawleri, the lattermost is identified as a non-cerapodan neornithis-
Emausaurus ernsti, Scelidosaurus harrisonii and Eurypoda chian in the derivative SRC tree: Fig. 4) that form a pectinate
(Ankylosauria + Stegosauria) is comparatively well- series of successively closer sister taxa to Cerapoda.
supported by total-evidence bootstrap proportions and Referral of Stormbergia to Neornithischia is supported
Bremer support (Fig. 2). The characters that unambiguously by only a single unambiguous character (possession of an
support the monophyly of this clade (Appendix 4) have obturator process on the ischium, Character 184, Appendix
already been identified by Sereno (1986, 1999a), with the 2), and measures of support for this node are weak (Bremer
exception of the presence of cortical remodelling of cra- support = +1, bootstrap = 33%). Referral of this taxon to
nial elements (Character 89, Appendix 2), which has been Neornithischia is, therefore, preliminary (although we have
previously considered to be limited to ankylosaurs, but is not yet been able to identify evidence that might support
also known in Scutellosaurus (UCMP 130580, jugal; Rosen- its referral to any other ornithischian clade) and a better
baum & Padian 2000: fig. 5C), Emausaurus (SGWG 85) and supported phylogenetic position must await the discovery of
Scelidosaurus (BMNH R1111; Carpenter 2001). specimens of this taxon with cranial material.
The phylogenetic position of Scelidosaurus harrisonii Apparent support is also weak for the position of Agil-
has proved controversial, despite being known from a num- isaurus and Hexinlusaurus as basal neornithischians. How-
ber of well-preserved specimens. This is probably because ever, bootstrap support for the node supporting the clade Oth-
no comprehensive description of the taxon has been car- nielia + Cerapoda, to the exclusion of Agilisaurus and Hexin-
ried out since the work of Owen (1861a, 1863), although lusaurus, is relatively high (79%) in the reduced bootstrap
a redescription is in progress (D.B.N. unpublished results). analysis (Fig. 4). Five unambiguous characters (Appendix 4)
Sereno (1986, 1999a) suggested 16 characters that link an- suggest that Hexinlusaurus is more closely related to cera-
kylosaurs and stegosaurs (as the clade Eurypoda) to the ex- podans than Agilisaurus. This is of interest as many authors
clusion of Scelidosaurus. Carpenter (2001) made the most have synonymised the two, although recent work (Barrett
extensive case to date in favour of positioning Scelidosaurus et al. 2005) supports generic-level distinction. Four addi-
as the most basal ankylosaur; however, Carpenter did not tional steps are required for Agilisaurus and Hexinlusaurus
include the eurypodan characters of Sereno (1999a) in his to group as sister taxa, although this does not represent a
analysis. At least four of the five synapomorphies identified significantly worse explanation of the data (Templeton test,
by Carpenter as linking Scelidosaurus with ankylosaurs ap- p = 0.10–0.35). Constraining these taxa as ornithopods (as
pear to have a more widespread distribution within Thyreo- suggested by previous authors, e.g. Weishampel & Heinrich
phora (Maidment et al., 2006; R. J. B. & S. C. R. Maidment, 1992; Norman et al. 2004c) requires 13 additional steps and
unpublished results). is a significantly worse explanation of the data than the most
This analysis does not find a sister group relationship parsimonious topology (Templeton test, p = 0.01–0.02).
between Scelidosaurus and ankylosaurs. A large number of The phylogenetic position of Othnielia is problematic.
anatomical features are shared by stegosaurs and ankylo- Othnielia has been invariably referred to Ornithopoda, usu-
saurs, but are absent in Scelidosaurus (Appendix 4; see also ally as a member of Hypsilophodontidae (e.g. Weishampel &
Sereno 1986, 1999a). It is possible that a number of these Heinrich 1992); by contrast, the derivative SRC tree (Fig. 4)
features could have evolved in parallel in both clades as ad- recovered by this analysis excludes Othnielia from Cerapoda,
aptations to large size (e.g. development of the preacetabular as a result of the retention of a relatively short postacetabular
process of the ilium, Character 166, Appendix 2; reduction process (Character 174, Appendix 2) and a relatively reduced
of the fourth trochanter, Character 201, Appendix 2), but and splint-like first metatarsal (Character 211, Appendix 2).
that this homoplasy has not yet been recognised as a result However, the position of Othnielia appears to be unstable
of the scarcity of well-preserved Early and Middle Jurassic and weakly supported and this may reflect the near-absence
eurypodans. It is also possible that the distribution of charac- of cranial data for this taxon.
ter states has been complicated by coding both Stegosauria
and Ankylosauria as supraspecific clades: a full considera-
tion of the phylogenetic position of Scelidosaurus requires
Ornithopoda
an analysis that includes a large number of stegosaur and A monophyletic Ornithopoda is not recovered in the SCC tree
ankylosaur taxa. Unfortunately such an analysis has not yet (Fig. 2), but is present in the derivative SRC tree (Fig. 4) fol-
been carried out and is beyond the scope of this work. How- lowing the a posteriori pruning of five unstable taxa, includ-
ever, the evidence presented by this analysis casts doubts ing the putative ornithopods Yandusaurus, Zephyrosaurus
upon interpretation of Scelidosaurus as a basal ankylosaur. and Talenkauen. However, the content of the ornithopod clade
Constraining Scelidosaurus to be the sister group of Ankylo- recovered in the derivative SRC tree (Fig. 4) differs signific-
sauria results in 2270 trees of 485 steps (8 extra steps), but is antly from that suggested by previous analyses. A number of
taxa generally referred to Ornithopoda (heterodontosaurids, alosaurs (Maryańska et al. 2004), the prepubic process of the
Agilisaurus, Hexinlusaurus, Othnielia) are here positioned pubis is elongated in some basal neornithischians (Hexin-
elsewhere within Ornithischia; this suggests that Ornitho- lusaurus, He & Cai 1984), pachycephalosaurs (Maryańska
poda, as conceived by some previous authors (e.g. Sereno et al. 2004: fig. 21.4C) and basal ceratopsians (e.g. Psit-
1986, 1999a; Norman et al. 2004c), may be polyphyletic; al- tacosauridae, Sereno 1987; Archaeoceratops, You & Dodson
ternatively, although we do not promote this view (we prefer 2003) and the obturator process is present on the ischium
the stem-based definition for Ornithopoda given in Table 1), in basal neornithischians (Stormbergia, SAM-PK-K1105,
it is conceivable that the definition of Ornithopoda could be BMNH R11000, Butler 2005; Agilisaurus, ZDM T6011;
expanded to include some, or all, of these taxa, as well as Hexinlusaurus, He & Cai 1984, ZDM T6001). Thus the ap-
ceratopsians and pachycephalosaurs. parently strong support for Ornithopoda found by previous
Hypsilophodontidae, as generally conceived analyses (e.g. Sereno 1999a) may be in part an artefact of an
(Weishampel & Heinrich 1992), does not form a mono- incorrect phylogenetic placement of heterodontosaurids.
phyletic clade; instead ‘hypsilophodontids’ appear to
represent a paraphyletic grade of basal neornithischian and
basal ornithopod taxa (see below). More derived ornithopod
Hypsilophodontidae
relationships generally follow that suggested by previous Early conceptions of Hypsilophodontidae were of an impli-
authors (e.g. Weishampel et al. 2003). citly paraphyletic ‘plexus’ of plesiomorphic, small, bipedal
The derivative SRC tree (Fig. 4) suggests that Ornitho- ornithischians (e.g. Thulborn 1971). Such a grouping was
poda includes a paraphyletic assemblage of ‘hypsilophodon- incompatible with cladistic study and a number of authors
tids’ (Orodromeus makelai, Jeholosaurus shangyuanensis, attempted to develop a more restrictive, monophyletic, defin-
Hypsilophodon foxii, Parksosaurus warreni, Gasparinisaura ition of the group (e.g. Norman 1984; Sereno 1984, 1986;
cincosaltensis, Bugenasaura infernalis, Thescelosaurus neg- Cooper 1985; Sues & Norman 1990; Weishampel & Heinrich
lectus) and Iguanodontia (comprising rhabdodontids, tenon- 1992). Sereno (1986) and Weishampel & Heinrich (1992)
tosaurs, dryosaurids and ankylopollexians). However, sup- considered the following taxa as members of Hypsilopho-
port is weak at the base of Ornithopoda, with most basal dontidae: Thescelosaurus neglectus, Othnielia rex, Hypsilo-
nodes having very low (less than 50%) total-evidence boot- phodon foxii, Zephyrosaurus schaffi, Orodromeus makelai
strap values. The characters that support basal nodes in the and Yandusaurus (the taxa Yandusaurus hongheensis and
derivative SRC tree tend to have limited or poorly understood Hexinlusaurus multidens were included as a single OTU;
distributions. For example, Ornithopoda is supported by the however, these taxa are distinguishable at generic level: Bar-
presence of a fossa-like depression on the premaxilla–maxilla rett et al. 2005). Sues & Norman (1990) referred a num-
boundary (Character 13, Appendix 2), the possession of nar- ber of other small ornithopods (e.g. Agilisaurus louderbacki,
row and elongate (more than twice as long as wide) frontals Gongbusaurus shiyii, ‘Gongbusaurus’ wucaiwanensis) to
that contact the nasals anterior to the orbits (Character 64, Hypsilophodontidae, while Bugenasaura infernalis has been
Appendix 2), and the possession of a foramen positioned referred to the clade by Galton (1999). However, recent ana-
on the dorsal part of the surangular–dentary joint (Charac- lyses (Scheetz 1998, 1999; Winkler et al. 1998; Weishampel
ter 105, Appendix 2). The first character is only known in et al. 2003; Norman et al. 2004c) have found little evidence
Orodromeus, Jeholosaurus and Hypsilophodon. It appears for hypsilophodontid monophyly.
to be absent in more derived ornithopods and a similar dis- A monophyletic Hypsilophodontidae was not recovered
tribution occurs for the second character (also present in by this analysis (Figs 2–4). Although ‘hypsilophodontids’ are
Zephyrosaurus, Sues 1980). generally thought of as rather anatomically conservative, it is
This analysis suggests that Ornithopoda is a poorly- clear that they are actually relatively diverse, with some taxa
supported clade and the weak support at the base of Or- (e.g. Thescelosaurus) showing greater similarities to more
nithopoda may be a result of the instability of fragment- derived, iguanodontian ornithopods, while other taxa appear
ary (e.g. Zephyrosaurus, Yandusaurus) or incompletely de- to have much more plesiomorphic morphologies (e.g. Oro-
scribed (e.g. Talenkauen) taxa. A further cause of the weak dromeus) and some may not even be referable to Ornithopoda
support for Ornithopoda may be the repositioning of het- (e.g. Agilisaurus, Hexinlusaurus, Othnielia).
erodontosaurids as basal ornithischians. Heterodontosaurids ‘Hypsilophodontids’ appear to represent a grade of
have effectively acted as an outgroup for analyses of ornitho- basal neornithischian and basal ornithopod taxa. Further
pod phylogeny. For instance, Sereno (1999a) suggested that work should be carried out on the relationships of these basal
the following characters were derived features of Euornitho- taxa. This analysis finds weak support for some groupings,
poda: reduction of the external antorbital fenestra; closure such as Gasparinisaura and Parksosaurus and more in depth
of the external mandibular fenestra; reduction of the delto- work in this part of the tree might clarify such possibilities.
pectoral crest of the humerus; enlargement of the prepubic
process of the pubis; presence of an obturator process on the
ischium. The plesiomorphic state for each of these characters
Marginocephalia
is present in Heterodontosauridae. However, if heterodonto- Only limited character evidence has been discovered pre-
saurids are repositioned basally within Ornithischia it be- viously for Marginocephalia (although see Xu et al. 2006),
comes apparent that each of the above characters suggested leading some authors to doubt its validity (Dodson 1990; Sul-
by Sereno (1999a) as diagnostic for Euornithopoda actually livan 2006). This analysis supports marginocephalian mono-
has a more widespread distribution within Cerapoda. For in- phyly; however, total-evidence bootstrap values (less than
stance, the external mandibular fenestra is closed in basal 50%) and Bremer support (+1) are very low for this clade.
neornithischians (e.g. Agilisaurus, Peng 1992), ceratopsians This might suggest that the clade is supported by a relatively
(e.g. Zhao et al. 1999; You & Dodson 2004) and pachyceph- low number of characters. However, character optimisation
(Appendix 4) indicates that Marginocephalia is supported by ceratopsian node (including Micropachycephalosaurus) is

8 unambiguous synapomorphies and as many as 17 char- +1. To test the effect of Micropachycephalosaurus and Sten-
acters under accelerated transformation (ACCTRAN). This opelix upon the apparent support for Ceratopsia an n-taxon
suggests that there is considerable character evidence for backbone constraint was defined which specified that Chaoy-
Marginocephalia, but that support for the clade is obscured angsaurus, Psittacosauridae and neoceratopsians were more
by one or more wildcard taxa. The putative basal pachyceph- closely related to each other than to ornithopods, pachyceph-
alosaur Stenopelix valdensis and the problematic cerapodan alosaurs, or non-cerapodans; however, it did not specify the
Micropachycephalosaurus hongtuyanensis are identified be- position of Micropachycephalosaurus, Stenopelix, or the five
low as wildcard taxa and it is possible that the apparent low wildcard taxa identified by the derivative SRC tree (Fig. 4).
support for Marginocephalia may result, in part, from the PAUP∗ was then used to search for the shortest trees in-
instability of these taxa. To test this possibility a backbone compatible with the backbone constraint. The shortest tree
constraint was written that did not specify the position of recovered was of 481 steps (Bremer support of +4), suggest-
Stenopelix, Micropachycephalosaurus, or the five unstable ing that strong support exists for the ceratopsian clade, but is
taxa identified in the derivative SRC tree (Fig. 4), but spe- obscured by the unstable nature of several taxa. To confirm
cified that all other pachycephalosaurs and ceratopsians were this, we pruned seven taxa (see above) including Micropachy-
more closely related to one another than any of them were cephalosaurus and Stenopelix from the set of trees recovered
to ornithopods, thyreophorans, or non-cerapodan neornithis- by the bootstrap analysis and generated a new majority-rule
chians. This constraint had a Bremer support of +4. This in- bootstrap reduced consensus tree, which demonstrates very
creased Bremer support suggests that there is relatively strong high (89%) bootstrap support for Ceratopsia.
‘hidden’ support for Marginocephalia. To confirm this, we Why does Micropachycephalosaurus act as a wildcard
pruned the five unstable taxa identified in the derivative SRC and why does this taxon group with ceratopsians when it
tree, Micropachycephalosaurus and Stenopelix from the set has been traditionally referred to Pachycephalosauria (Dong
of trees recovered by the bootstrap analysis and generated 1978; Maryańska et al. 2004)? The holotype of Micropachy-
a new 50% majority-rule bootstrap reduced consensus tree, cephalosaurus (IVPP V 5542), originally described by Dong
which demonstrates increased (70%) bootstrap support for (1978), is extremely fragmentary and comprises a quadrate
Marginocephalia. and partial tooth row, vertebral elements (including an in-
Three key characters have been identified by previous complete sacrum), a partial ilium and the partial left femur
authors (Sereno 1984, 1986, 1987, 1999a, 2000; Cooper and tibia. Dong (1978) referred Micropachycephalosaurus to
1985; Maryańska & Osmólska 1985) as supporting mar- Pachycephalosauria on the basis of a supposedly thickened
ginocephalian monophyly: posterior expansion of the pari- skull roof; however, no material of the skull roof is cur-
etosquamosal obscures the occiput in dorsal view, premaxil- rently present in the holotype specimen (R. J. B. pers. obs.,
lae excluded from the border of the internal nares, postpubis 2004; R. J. B. & Q. Zhao, unpublished results). In addition,
reduced or absent (Characters 68, 84, 188/189, Appendix other pachycephalosaurian synapomorphies (e.g. preacetab-
2). Other suggested marginocephalian synapomorphies (see, ular process of ilium expands mediolaterally anteriorly in
e.g., Cooper 1985; Sereno 1987) have not been supported by dorsal view; elongate caudal ribs) are absent. Cerapodan fea-
subsequent work. This analysis supports the validity of these tures (particularly of the femur) are present, but few charac-
three marginocephalian characters. ters can be identified that unambiguously link Micropachy-
A number of other potential marginocephalian synapo- cephalosaurus with pachycephalosaurs, ornithopods or cer-
morphies are identified here. Unambiguous synapomorphies atopsians. The only character that can be identified as po-
include: absence of elliptic fossa on the nasals (although a tentially linking Micropachycephalosaurus with ceratopsi-
fossa is present in at least one basal ceratopsian, Liaoceratops ans is the enlargement of the lateral condyle of the quadrate
yanzigouensis), three premaxillary teeth, elongate and strap- (Character 63, Appendix 2); however, this character is ho-
like scapula blade and the absence of an obturator process moplastic, with the derived state also known in heterodonto-
on the ischium (Characters 19, 112, 150, 184, Appendix 2). saurids (Heterodontosaurus, SAM-PK-K337) and some or-
ACCTRAN indicates additional potential synapomorphies, nithopods (e.g. Weishampel et al. 2003). Only one additional
most of which have rather uncertain distributions and suffer step is required for Micropachycephalosaurus to group with
from missing data. For example, node-like ornamentation either pachycephalosaurs or ornithopods, instead of ceratop-
is present on the jugal–postorbital bar and angular (Charac- sians. A ceratopsian identity appears to be poorly supported
ters 41, 108, Appendix 2) of pachycephalosaurs and the basal and the position of Micropachycephalosaurus within Cera-
ceratopsians Chaoyangsaurus youngi and Archaeoceratops poda cannot be clarified with any certainty (R. J. B. & Q.
oshimai. However, the distribution of this character remains Zhao, unpublished results).
uncertain as ornamentation is absent in other basal ceratop-
sians such as Psittacosauridae.
Pachycephalosauria
Pachycephalosauria is a well-established clade and all pub-
Ceratopsia
lished ornithischian phylogenetic analyses have found it to
Ceratopsia is recovered as a monophyletic clade in this ana- be monophyletic. This analysis finds Pachycephalosauria
lysis, although decay indices and bootstrap values suggest to be a monophyletic clade including Stenopelix valdensis,
that support for the clade is weak. However, this may result Wannanosaurus yansiensis, Goyocephale lattimorei, Homa-
from the inclusion within the clade of the fragmentary and locephale calathocercos and Pachycephalosauridae; how-
problematic taxon Micropachycephalosaurus hongtuyanen- ever, Bremer support (+1) and total-evidence bootstrap val-
sis, which appears to act as a ‘wildcard’, or the unstable ues (55%) show surprisingly low support for Pachyceph-
position of the putative basal pachycephalosaur Stenopelix alosauria (Fig. 2), although bootstrap support is stronger
valdensis (see below), or both. The Bremer support for the for more derived groupings within the clade. However, as
with Ceratopsia, the real support for the clade may be ob- Ornithischia: basal ornithopod phylogeny. Removing hetero-
scured by the presence of one or more ‘wildcard’ taxa. The dontosaurids here results in a number of taxa long considered
most obvious candidate is the problematic basal taxon Sten- as ornithopods (e.g. Agilisaurus louderbacki, Hexinlusaurus
opelix. To test this possibility, a backbone constraint was multidens, Othnielia rex) grouping outside of Cerapoda, as
defined that specified that Wannanosaurus, Goyocephale, non-ornithopods. This analysis suggests that the number of
Homalocephale and Pachycephalosauridae are more closely taxa that can be referred to Ornithopoda is more restrictive
related to one another than to ceratopsians, ornithopods, or than hypothesised by previous workers. In addition, recog-
non-cerapodans (the phylogenetic positions of the five wild- nising robust ornithopod synapomorphies is extremely diffi-
card taxa identified by reduced consensus, Stenopelix and cult and basal relationships (amongst the ‘hypsilophodontid’
the problematic taxon Micropachycephalosaurus were not taxa) are poorly resolved and weakly supported. Consider-
specified by this constraint). The shortest tree incompatible able future work is required to determine exactly which taxa
with this constraint was 480 steps long (3 steps longer than can be referred with confidence to this clade, what charac-
the MPTs), suggesting that unstable taxa are decreasing the ters diagnose the clade and how basal taxa are related to one
apparent support for Pachycephalosauria. A new majority- another.
rule bootstrap reduced consensus tree, following the prun-
ing of seven taxa (see above) demonstrates exceptionally
high (95%) bootstrap support for Pachycephalosauria (Wan- Implications for phylogenetic taxonomy of
nanosaurus, Goyocephale, Homalocephale, Pachycephalo- Ornithischia
sauridae). Thus, although support for pachycephalosaurian Sereno (1998, 1999b) provided node-based and stem-based
monophyly may be strong, it is obscured by the presence phylogenetic definitions for a large number of ornithischian
in the data set of a number of wildcard taxa of uncertain clades; alternative definitions for a number of major ornithis-
position. chian clades have since been provided by Buchholz (2002),
Wagner (2004), various authors in Weishampel et al. (2004a)
and this study (Table 1). The results of this analysis highlight
some of the problems that can occur when clade names are
given phylogenetic definitions on the basis of assumed stabil-
Conclusions ity and later phylogenetic analyses generate alternate topolo-
gies. For instance, this analysis positions heterodontosaurids
Stability, instability, and future directions
near to the base of Ornithischia, rather than as ornithopods as
in ornithischian phylogeny suggested by many previous authors. Sereno (1998) provided
The results of this phylogenetic analysis differ in many re- a node-based definition for the clade Ornithopoda: ‘Hetero-
spects from those of previous analyses and reinforce the dontosaurus, Parasaurolophus, their most recent common
importance of constant reassessment of phylogenetic hypo- ancestor and all descendents’. When this definition is ap-
theses. Furthermore, the results of this analysis highlight plied to the phylogeny produced by this analysis (Fig. 4),
areas of stability within ornithischian phylogeny and also virtually all ornithischians (with the exception of Pisano-
problematic areas in which future efforts should be concen- saurus mertii), including ankylosaurs, stegosaurs, ceratop-
trated. sians and pachycephalosaurs, are included within Ornitho-
At a broad-scale, the widely accepted framework of or- poda. This is a radically different, and rather unsatisfactory,
nithischian phylogeny, developed by Norman (1984), Sereno taxonomic content to that currently understood. The prob-
(1984, 1986, 1991a, 1999a), Cooper (1985) and Maryańska lem is avoided if a conservative stem-based definition is used
& Osmólska (1985), amongst others, has proved relatively for Ornithopoda (see Table 1; Buchholz 2002; Wagner 2004;
stable (Fig. 4). There appears to be a basal split within Ornith- Norman et al. 2004c). Likewise, Sereno (1998) provided a
ischia between thyreophorans and neornithischians. Within stem-based definition for the clade Euornithopoda: ‘All or-
Neornithischia two monophyletic clades occur, Ornithopoda nithopods closer to Parasaurolophus than to Heterodonto-
and Marginocephalia; the latter is subdivided into Ceratop- saurus.’ Applying this definition to the phylogeny produced
sia and Pachycephalosauria. In many cases this analysis has by this analysis results in Euornithopoda effectively refer-
strengthened this basic framework; for example, by identify- ring to the same clade as Genasauria. Again, this is an un-
ing additional marginocephalian synapomorphies. However, satisfactory taxonomic content for Euornithopoda, radically
a number of unstable and problematic areas within this frame- different from its currently understood content. In this case it
work have also been identified. would probably have been better not to provide a phylogen-
Heterodontosaurids have long been recognised as the etic definition for Euornithopoda, given the known unstable
most unstable of ornithischian taxa and the clade has been phylogenetic position of the external specifier, Heterodonto-
positioned in four very different positions within Ornith- saurus tucki.
ischia. This analysis positions heterodontosaurids as non- Another taxonomic problem is highlighted by this ana-
genasaurian basal ornithischians, but we additionally note lysis. Iguanodontia has generally been used for those taxa
that recent authors have found anatomical evidence and more derived within Ornithopoda than Hypsilophodontidae.
phylogenetic support for the idea that heterodontosaurids However, such a definition is problematic if, as sugges-
may form the sister clade to Marginocephalia (e.g. Xu et al. ted by this analysis, Hypsilophodontidae is paraphyletic.
2006). Resolving the position of heterodontosaurids within Sereno (1998) proposed the stem-based definition, ‘All eu-
Ornithischia is one of the most important tasks facing ornith- ornithopods closer to Parasaurolophus than to Hypsilopho-
ischian phylogeneticists and future work should aim to com- don’. Application of this definition to the phylogeny pro-
bine the data set provided here with that of Xu et al. (2006). duced by this analysis suggests that taxa such as Thescelo-
The removal of heterodontosaurids from the base of Or- saurus neglectus and Parksosaurus warreni are iguanodon-
nithopoda highlights another major area of instability within tians. However, such taxa have generally been considered as
‘hypsilophodontid’ basal ornithopods, outside of Iguanodon- — 2001. Tooth wear and possible jaw action of Scelidosaurus harrisonii
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H. Osmólska (eds) The Dinosauria, Second Edition. University of transitional features from the Late Jurassic of northwestern China.
California Press, Berkeley. Proceedings of the Royal Society B 273: 2135–2140.
—, Makovicky, P. J., Wang, X.-L., Norell, M. A. & You, H.-L. 2002. A Bugenasaura infernalis Galton, 1995: Galton (1995,
ceratopsian dinosaur from China and the early evolution of Ceratopsia. 1999); SDSM 7210 (holotype), MOR 979.
Nature 416: 314–317.
Chaoyangsaurus youngi Zhao et al., 1999: Zhao et al.
Yates, A. M. 2003. A new species of the primitive dinosaur Thecodon-
tosaurus (Saurischia: Sauropodomorpha) and its implications for the (1999); IGCAGS V371 (holotype).
systematics of early dinosaurs. Journal of Systematic Palaeontology 1: Dryosauridae Milner & Norman, 1984: Janensch (1955),
1–42. Galton (1981, 1983, 1989), Galton & Taquet (1982);
You, H.-L. & Dodson, P. 2003. Redescription of neoceratopsian dinosaur BMNH R185, R186, holotype of Valdosaurus canalicu-
Archaeoceratops and early evolution of Neoceratopsia. Acta Palaeon-
latus; MB, numerous cranial and postcranial specimens
tologica Polonica 48: 261–272.
— & — 2004. Basal Ceratopsia. Pp. 478–493 in D. B. Weishampel, P. of Dysalotosaurus lettowvorbecki (see Galton 1981,
Dodson & H. Osmólska (eds) The Dinosauria, 2nd ed. University of 1983).
California Press, Berkeley. Echinodon becklesii Owen, 1861b: Galton (1978), Nor-
—, Xu, X. & Wang, X.-L. 2003. A new genus of Psittacosauridae (Di- man & Barrett (2002); BMNH 48209, 48210 (lecto-
nosauria: Ornithopoda) and the origin and early evolution of margino-
types); BMNH 48211–48215 (paralectotypes).
cephalian dinosaurs. Acta Geologica Sinica 77: 15–20.
Zhao, X.-J., Cheng, Z.-W. & Xu, X. 1999. The earliest ceratopsian from Emausaurus ernsti Haubold, 1990: Haubold (1990), Nor-
the Tuchengzi Formation of Liaoning, China. Journal of Vertebrate man et al. (2004b); SGWG 85 (holotype).
Paleontology 19: 681–691. Gasparinisaura cincosaltensis Coria & Salgado 1996:
Coria & Salgado (1996), Salgado et al. (1997).
Goyocephale lattimorei Perle et al., 1982: Perle et al.
(1982), Sereno (2000), Maryańska et al. (2004).
Heterodontosaurus tucki Crompton & Charig, 1962:
Appendix 1: Specimens and Crompton & Charig (1962), Santa Luca et al. (1976),
references used for coding Santa Luca (1980), Weishampel & Witmer (1990b),
Norman et al. (2004c); SAM-PK-K337 (holotype);
operational taxonomic units SAM-PK-K1332 (referred specimen).
Euparkeria capensis Broom, 1913: Ewer (1965), Gower Hexinlusaurus multidens (He & Cai, 1983): He & Cai
& Weber (1998). (1984), Barrett et al. (2005); ZDM T6001 (holotype).
Marasuchus liloensis (Romer, 1972): Bonaparte (1975), Homalocephale calathocercos Maryańska & Osmólska,
Sereno & Arcucci (1994). 1974: Maryańska & Osmólska (1974), Maryańska
Herrerasaurus ischigualastensis Reig, 1963: Novas (1990), Sereno (2000), Maryańska et al. (2004).
(1993), Sereno (1993), Sereno & Novas (1993); MCZ Hypsilophodon foxii Huxley, 1869: Galton (1974a);
7063, 7064. BMNH R197 (holotype); other BMNH specimens (see
Abrictosaurus consors (Thulborn, 1974): Thulborn Galton 1974a).
(1974), Hopson (1975); BMNH RUB54 (holotype). Jeholosaurus shangyuanensis Xu et al., 2000: Xu et al.
Agilisaurus louderbacki Peng, 1990: Peng (1992, 1997), (2000); IVPP V12529 (holotype); IVPP V12530.
Barrett et al. (2005); ZDM 6011 (holotype). Lesothosaurus diagnosticus Galton, 1978: Thulborn
Anasibetia saldiviai Coria & Calvo, 2002: Coria & Calvo (1970a, 1972), Santa Luca (1984), Weishampel &
(2002). Witmer (1990a), Sereno (1991a), Norman et al. (2004a),
Ankylopollexia Sereno, 1986: Gilmore (1909), Galton Butler (2005); BMNH RUB17, RUB23 (syntypes);
& Powell (1980), Norman (1980, 1986, 2002, 2004), BMNH R11956, R8501 (referred specimens); SAM-PK-
Norman & Weishampel (1990); OUM J330 (holotype, K400, K401, K1106 (referred specimens).
‘Cumnoria’ prestwichii); BMNH, numerous specimens Liaoceratops yanzigouensis Xu et al., 2002: Xu et al.
referable to Iguanodon spp. (2002); IVPP V12738 (holotype).
Ankylosauria Osborn, 1923: Ostrom (1970), Maryańska Lycorhinus angustidens Haughton, 1924: Gow (1975,
(1977), Coombs (1978), Coombs & Maryańska (1990), 1990), Hopson (1975, 1980); SAM-PK-K3606
Lee (1996), Carpenter (2001, 2004), Carpenter et al. (holotype); BP/1/4244, BP/1/5253 (referred, Gow
(2001), Xu et al. (2001), Vickaryous et al. (2001, 1990).
2004); BMNH R175, Polacanthus foxii (holotype); Micropachycephalosaurus hongtuyanensis Dong, 1978:
BYU, material of Gastonia burgei (many specimens); Dong (1978); IVPP V5542 (holotype).
CEUM, holotypes and referred material of Gastonia Othnielia rex (Marsh, 1877b): Galton & Jensen (1973),
burgei, Animantarx ramaljonesi, Cedarpelta bilbeyhal- Galton (1983); BYU ESM-163R.
lorum (see Kirkland 1998; Carpenter et al. 1999, 2001); Orodromeus makelai Horner & Weishampel, 1988:
MWC, holotype and referred material of Mymoorapelta Horner & Weishampel (1988), Scheetz (1999); MOR
maysi (see Kirkland & Carpenter 1994; Kirkland et 294 (holotype), MOR 403, 473, 623, 1141 (referred spe-
al. 1998); IVPP V12560 (holotype of Liaoningosaurus cimens).
paradoxus). Pachycephalosauridae Sternberg, 1945: Brown & Sch-
Archaeoceratops oshimai Dong & Azuma, 1997: Dong laikjer (1943), Maryańska & Osmólska (1974), Sues
& Azuma (1997), You & Dodson (2003); IVPP V11114 & Galton (1987), Maryańska (1990), Sereno (2000),
(holotype); IVPP V11115 (paratype). Maryańska et al. (2004); BMNH R10055 (cast of
holotype skull of Stegoceras), ZPAL MgD-I/104 (holo- Wannanosaurus yansiensis Hou, 1977: Hou (1977), Ser-
type of Prenocephale prenes). eno (2000); IVPP V4447 (holotype), IVPP V4447.1
Parksosaurus warreni (Parks, 1926): Parks (1926), Stern- (paratype).
berg (1940), Galton (1973b). Yandusaurus hongheensis He, 1979: He & Cai (1984);
Pisanosaurus mertii Casamiquela, 1967: Casamiquela GCC V20501 (holotype).
(1967), Bonaparte (1976), Weishampel & Witmer Zephyrosaurus schaffi Sues, 1980: Sues (1980); MCZ
(1990a), Sereno (1991a), Báez & Marsicano (2001), 4392 (holotype).
Norman et al. (2004a).
Psittacosauridae Osborn, 1923: Sereno (1987, 1990a, b,
2000), Sereno & Chao (1988), Sereno et al. (1988), You
et al. (2003); IVPP V12704 (Hongshanosaurus holo- Appendix 2: Character list
type); IVPP V738, 740–1, 749 (Psittacosaurus sinensis The following references were used for compiling previously
holotype and referred specimens); IVPP V7705 (Psit- used characters: Maryańska & Osmólska (1984, 1985), Mil-
tacosaurus meileyingensis holotype). ner & Norman (1984), Norman (1984, 1990, 1998, 2004),
Rhabdodontidae Weishampel et al., 2003: Pincemaille- Sereno (1984, 1986, 1987, 1999a, 2000), Cooper (1985),
Quillévéré (2002), Weishampel et al. (2003); BMNH Gauthier (1986), Forster (1990), Sereno & Dong (1992),
specimens of Zalmoxes robustus (see Weishampel et al. Weishampel & Heinrich (1992), Novas (1993, 1996), Ser-
2003 for full list). eno & Novas (1993), Coria & Salgado (1996), Winkler et al.
Scelidosaurus harrisonii Owen, 1861a: Owen (1861a, (1997), Zhao et al. (1999), Carpenter (2001), Makovicky
1863), Norman et al. (2004b); BMNH R1111 (lectotype); (2001), Williamson & Carr (2002), Xu et al. (2002), Rauhut
(2003), Yates (2003), Weishampel et al. (2003), You et al.
BMNH R5909, R6704; BRSMG Ce12785; CAMSM X
(2003), Langer (2004), Norman et al. (2004b, c), Novas
39256. et al. (2004), Butler (2005).
Scutellosaurus lawleri Colbert, 1981: Colbert (1981),
Coombs et al. (1990), Sereno (1991a), Rosenbaum & 1. Skull proportions: 0, preorbital skull length more than
Padian (2000), Norman et al. (2004b); MNA P1.175 45% of basal skull length; 1, preorbital length less than
(holotype); MNA P1.1752 (paratype); UCMP 130580, 40% of basal skull length.
130581, 170829, 175166, 175167, 175168 (referred spe- 2. Skull length (rostral–quadrate): 0, 15% or less of body
cimens). length; 1, 20–30% of body length.
Stegosauria Marsh, 1877a: Gilmore (1914), Galton 3. Neomorphic rostral bone, anterior to premaxilla: 0, ab-
sent; 1, present.
(1982a, b, 1985, 1988, 1990, 1991), Dong et al. (1983),
4. Rostral bone, anteriorly keeled and ventrally pointed:
Sereno & Dong (1992), Galton & Upchurch (2004b);
0, absent; 1, present.
Cast of IVPP V6728 (holotype skull of Huayango- 5. Rostral bone, ventrolateral processes: 0, rudimentary;
saurus taibaii); ZDM 7001 (Huayangosaurus taibaii); 1, well-developed.
BMNH 46013, R5902 (holotype and referred specimen, 6. Premaxilla, edentulous anterior region: 0, absent, first
Dacentrurus armatus); BMNH R1989, R3167 (holo- premaxillary tooth is positioned adjacent to the sym-
type and referred specimen, Lexovisaurus durobriven- physis; 1, present, first premaxillary tooth is inset the
sis); MB, Kentrosaurus aethiopicus syntypes and re- width of one or more crowns.
ferred material (see Galton 1982b, 1988); IVPP V2300, 7. Premaxilla, posterolateral process, length: 0, does not
CV 202, 203 (all Chialingosaurus kuani); CV 205, 206 contact lacrimal; 1, contacts the lacrimal, excludes
(Chungkingosaurus jiangbeiensis); CV 209, 210 (Tuoji- maxilla–nasal contact.
8. Oral margin of the premaxilla: 0, narial portion of the
angosaurus multispinus).
body of the premaxilla slopes steeply from the external
Stenopelix valdensis Meyer, 1857: Sues & Galton (1982),
naris to the oral margin; 1, ventral premaxilla flares
Sereno (1987); GZG 741/2 (holotype). laterally to form a partial floor of the narial fossa.
Stormbergia dangershoeki Butler, 2005: Butler (2005); 9. Position of the ventral (oral) margin of the premax-
SAM-PK-K1105 (holotype); BMNH R11000 (para- illa: 0, level with the maxillary tooth row; 1, deflected
type); BP/1/4885 (referred). ventral to maxillary tooth row.
Talenkauen santacrucensis Novas et al., 2004: Novas 10. Premaxillary foramen: 0, absent; 1, present.
et al. (2004). 11. Premaxillary palate: 0, strongly arched, forming a deep,
Tenontosaurus tilletti Ostrom, 1970: Ostrom (1970), For- concave palate; 1, horizontal or only gently arched.
ster (1990). 12. Overlap of the dorsal process of the premaxilla onto
Tenontosaurus dossi Winkler et al., 1997: Winkler et al. the nasal: 0, present; 1, absent.
13. Fossa-like depression positioned on the premaxilla–
(1997).
maxilla boundary: 0, absent; 1, present.
Thescelosaurus neglectus Gilmore, 1913: Gilmore
14. Premaxilla–maxilla diastema: 0, absent, maxillary
(1915), Sternberg (1940), Galton (1974b, 1995, 1997). teeth continue to anterior end of maxilla; 1, present,
Unnamed taxon (Coronosauria + Leptoceratopsidae): substantial diastema of at least one crown’s length
Brown & Schlaikjer (1940, 1942), Maryańska & between maxillary and premaxillary teeth.
Osmólska (1975), Sereno (2000), You & Dodson (2004); 15. Form of diastema; 0, flat; 1, arched ‘subnarial gap’
MOR 542 (Montanoceratops, referred specimen). between the premaxilla and maxilla.
16. Narial fossa surrounding external nares on lateral 43. Jugal–postorbital joint: 0, elongate scarf joint; 1, short
surface of premaxilla, position of ventral margin of butt joint.
fossa relative to the ventral margin of the premaxilla: 44. Jugal, form of postorbital process: 0, not expanded
0, closely approaches the ventral margin of the premax- dorsally; 1, dorsal portion of postorbital process is ex-
illa; 1, separated by a broad flat margin from the ventral panded posteriorly.
margin of the premaxilla. 45. Jugal–squamosal contact above infratemporal fenestra:
17. External nares, position of the ventral margin: 0, below 0, absent; 1, present.
the ventral margin of the orbits; 1, above the ventral 46. Jugal posterior ramus, forked: 0, absent; 1, present.
margin of the orbits. 47. Jugal, posterior ramus: 0, forms anterior and ventral
18. External naris size: 0, small, entirely overlies the margin of infratemporal fenestra; 1, forms part of pos-
premaxilla; 1, enlarged, extends posteriorly to overlie terior margin, expands towards squamosal.
the maxilla. 48. Jugal–quadratojugal contact: 0, overlapping; 1, tongue-
19. Deep elliptic fossa present along sutural line of the and-groove.
nasals: 0, absent; 1, present. 49. Postorbital, orbital margin: 0, relatively smooth curve;
20. Internal antorbital fenestra size: 0, large, generally at 1, prominent and distinct projection into orbit.
least 15% of the skull length; 1, very much reduced, 50. Postorbital: 0, T-shaped; 1, triangular and plate-like.
less than 10% of skull length, or absent. 51. Postorbital–parietal contact: 0, absent, or very narrow;
21. External antorbital fenestra: 0, present; 1, absent. 1, broad.
22. External antorbital fenestra, shape: 0, triangular; 1, oval 52. Contact between dorsal process of quadratojugal and
or circular. descending process of the squamosal: 0, present;
23. Additional opening(s) anteriorly within the antorbital 1, absent.
fossa: 0, absent; 1, present. 53. Quadratojugal, shape: 0, L-shaped, with elongate an-
24. Maxilla, prominent anterolateral boss articulates with terior process; 1, subrectangular with long axis vertical,
the medial premaxilla: 0, absent; 1, present. short, deep anterior process.
25. Maxilla, accessory anterior process: 0, absent; 54. Quadratojugal, ventral margin: 0, approaches the
1, present. mandibular condyle of the quadrate; 1, well-removed
26. Maxilla, buccal emargination: 0, absent; 1, present. from the mandibular condyle of the quadrate.
27. Eminence on the rim of the buccal emargination of 55. Quadratojugal, orientation: 0, faces laterally; 1, faces
the maxilla near the junction with the jugal: 0, absent; posterolaterally.
1, present. 56. Quadratojugal, transverse width: 0, mediolaterally
28. Slot in maxilla for lacrimal: 0, absent; 1, present. flattened; 1, transversely expanded and triangular in
29. Accessory ossification(s) in the orbit (palpeb- coronal section.
ral/supraorbital): 0, absent; 1, present. 57. Prominent oval fossa on pterygoid ramus of quadrate:
30. Palpebral/supraorbital: 0, free, projects into orbit from 0, absent; 1, present.
contact with lacrimal/prefrontal; 1, incorporated into 58. Quadrate lateral ramus: 0, present; 1, absent.
orbital margin. 59. Quadrate shaft: 0, anteriorly convex in lateral view;
31. Palpebral, shape in dorsal view: 0, rod-shaped; 1, plate- 1, reduced in anteroposterior width and straight in lat-
like with wide base. eral view.
32. Palpebral/supraorbital, number: 0, one; 1, two; 2, three. 60. Paraquadratic foramen or notch, size: 0, absent or
33. Free palpebral, length, relative to anteroposterior width small, opens between quadratojugal and quadrate;
of orbit: 0, does not traverse entire width of orbit; 1, large.
1, traverses entire width of orbit. 61. Paraquadratic foramen, orientation: 0, posterolateral
34. Exclusion of the jugal from the posteroventral margin aspect of quadrate shaft; 1, lateral aspect of quadrate
of the external antorbital fenestra by lacrimal–maxilla or quadratojugal.
contact: 0, absent; 1, present. 62. Paraquadratic foramen, position: 0, on quadrate-
35. Anterior ramus of jugal, proportions: 0, deeper than quadratojugal boundary; 1, located within quadrato-
wide, but not as deep as the posterior ramus of the jugal.
jugal; 1, wider than deep; 2, deeper than the posterior 63. Quadrate mandibular articulation: 0, quadrate con-
ramus of the jugal. dyles subequal in size; 1, medial condyle is larger
36. Widening of the skull across the jugals, chord from than lateral condyle; 2, lateral condyle is larger than
frontal orbital margin to extremity of jugal is more medial.
than minimum interorbital width: 0, absent; 1, present, 64. Paired frontals: 0, short and broad; 1, narrow and elong-
skull has a triangular shape in dorsal view. ate (more than twice as long as wide).
37. Position of maximum widening of the skull: 0, beneath 65. Supratemporal fenestrae: 0, open; 1, closed.
the jugal–postorbital bar; 1, posteriorly, beneath the 66. Supratemporal fenestrae, anteroposteriorly elongated:
infratemporal fenestra. 0, absent, fenestrae are subcircular to oval in shape;
38. Jugal (or jugal–epijugal) ridge dividing the lateral sur- 1, present.
face of the jugal into two planes: 0, absent; 1, present. 67. Parietal septum, form: 0, narrow and smooth; 1, broad
39. Epijugal: 0, absent; 1, present. and rugose.
40. Jugal boss: 0, absent; 1, present. 68. Parietosquamosal shelf: 0, absent; 1, present.
41. Node-like ornamentation on jugal, mostly on, or ventral 69. Parietosquamosal shelf, extended posteriorly as distinct
to, the jugal–postorbital bar: 0, absent; 1, present. frill: 0, absent; 1, present.
42. Jugal–postorbital bar, width broader than laterotem- 70. Composition of the posterior margin of the pari-
poral fenestra: 0, absent; 1, present. etosquamosal shelf: 0, parietal contributes only a small
portion to the posterior margin;1. parietal makes up at 99. Dorsal and ventral margins of the dentary: 0, converge
least 50% of the posterior margin. anteriorly; 1, subparallel.
71. Postorbital–squamosal bar: 0, bar-shaped; 1, broad, 100. Ventral flange on dentary: 0, absent; 1, present.
flattened. 101. Coronoid process: 0, absent or weak, posterodorsally
72. Postorbital–squamosal tubercle row: 0, absent; oblique, depth of mandible at coronoid is less than
1, present. 140% depth of mandible beneath tooth row; 1, well-
73. Enlarged tubercle row on the posterior squamosal: developed, distinctly elevated, depth of mandible at
0, absent; 1, present. coronoid is more than 180% depth of mandible beneath
74. Frontal and parietal dorsoventral thickness: 0, thin; tooth row.
1, thick. 102. Anterodorsal margin of coronoid process formed by
75. Paroccipital processes: 0, extend laterally and are posterodorsal process of dentary: 0, absent; 1, present.
slightly expanded distally; 1, distal end pendent and 103. Coronoid process, position: 0, posterior to dentition;
ventrally extending. 1, lateral to dentition.
76. Paroccipital processes, proportions: 0, short and deep 104. External mandibular fenestra, situated on dentary-
(height ≥ 1/2 length); 1, elongate and narrow. surangular-angular boundary: 0, present; 1, absent.
77. Posttemporal foramen/fossa, position: 0, totally en- 105. Small fenestra positioned dorsally on the surangular-
closed with the paroccipital process; 1, forms a notch in dentary joint: 0, absent; 1, present.
the dorsal margin of the paroccipital process, enclosed 106. Ridge or process on lateral surface of surangular, an-
dorsally by the squamosal. terior to jaw suture: 0, absent; 1, present, strong antero-
78. Supraoccipital, contribution to dorsal margin of fo- posteriorly extended ridge; 2, present, dorsally directed
ramen magnum: 0, forms entire dorsal margin of finger-like process.
foramen magnum; 1, exoccipital with medial pro- 107. Retroarticular process: 0, elongate; 1, rudimentary or
cess that restricts the contribution of the supraoccipi- absent.
tal. 108. Node-like ornamentation of the dentary and angular:
79. Basioccipital, contribution to the border of the foramen 0, absent; 1, present.
magnum: 0, present; 1, absent, excluded by exoccipit- 109. Level of jaw joint: 0, level with tooth row, or weakly
als. depressed ventrally; 1, strongly depressed ventrally,
80. Basisphenoid: 0, longer than, or subequal in length to, more than 40% of the height of the quadrate is below
basioccipital; 1, shorter than basioccipital. the level of the maxilla.
81. Prootic–basisphenoid plate: 0, absent; 1, present. 110. Mandibular osteoderm: 0, absent; 1, present.
82. Basal tubera, shape: 0, knob-shaped; 1, plate-shaped. 111. Premaxillary teeth: 0, present; 1, absent, premaxilla
83. Basipterygoid processes, orientation: 0, anteroventral; edentulous.
1, ventral; 2, posteroventral. 112. Premaxillary teeth, number: 0, six; 1, five; 2. four;
84. Premaxilla–vomeral contact: 0, present; 1, absent, ex- 3. three; 4. two.
cluded by midline contact between maxillae. 113. Premaxillary teeth, crown expanded above root:
85. Dorsoventrally deep (deeper than 50% of snout depth) 0, crown is unexpanded mesiodistally above root, no
median palatal keel formed of the vomers, pterygoids distinction between root and crown is observable;
and palatines: 0, absent; 1, present. 1, crown is at least moderately expanded above root.
86. Pterygovomerine keel, length: 0, less than 50% of pal- 114. Premaxillary teeth increase in size posteriorly: 0, ab-
ate length; 1, more than 50% of palate length. sent, all premaxillary teeth subequal in size; 1, present,
87. Pterygoid–maxilla contact at posterior end of tooth posterior premaxillary teeth are significantly larger in
row: 0, absent; 1, present. size than anterior teeth.
88. Pterygoquadrate rami, posterior projection of ventral 115. Maxillary and dentary crowns, shape: 0, apicobas-
margin: 0, weak; 1, pronounced. ally tall and blade-like; 1, apicobasally short and sub-
89. Cortical remodeling of surface of skull dermal bone: triangular; 2, diamond-shaped.
0, absent; 1, present. 116. Maxillary/dentary teeth, marginal ornamentations:
90. Predentary: 0, absent; 1, present. 0, fine serrations set at right angles to the margin of the
91. Predentary size: 0, short, posterior premaxillary teeth tooth; 1, coarse serrations (denticles) angle upwards at
oppose anterior dentary teeth; 1, roughly equal in length 45◦ from the margin of the tooth.
to the premaxilla, premaxillary teeth only oppose pre- 117. Enamel on maxillary/dentary teeth: 0, symmetrical;
dentary. 1, asymmetrical.
92. Predentary, rostral end in dorsal view: 0, rounded; 118. Apicobasally extending ridges on maxillary/dentary
1, pointed. teeth: 0, absent; 1, present.
93. Predentary, oral margin: 0, relatively smooth; 119. Apicobasally extending ridges on lingual/labial sur-
1, denticulate. faces of maxillary/dentary crowns confluent with mar-
94. Tip of predentary in lateral view: 0, does not project ginal denticles: 0, absent; 1, present.
above the main body of predentary; 1, strongly up- 120. Prominent primary ridge on labial side of maxillary
turned relative to main body of predentary. teeth: 0, absent; 1, present.
95. Predentary, ventral process: 0, single; 1, bilobate. 121. Prominent primary ridge on lingual side of dentary
96. Predentary, ventral process: 0, present, well-developed; teeth: 0, absent; 1, present.
1, very reduced or absent. 122. Position of maxillary/dentary primary ridge: 0, centre
97. Dentary symphysis: 0, V-shaped; 1, spout shaped. of the crown surface, giving the crown a relatively sym-
98. Dentary tooth row (and edentulous anterior portion) in metrical shape in lingual/labial view; 1, offset, giving
lateral view: 0, straight; 1, anterior end downturned. crown asymmetrical appearance.
123. At least moderately developed labiolingual expan- imum width; 1, elongate and strap-like, length is at
sion of crown (‘cingulum’) on maxillary/dentary teeth: least 9 times the minimum width.
0, present; 1, absent. 151. Scapula acromion shape: 0, weakly developed or ab-
124. Heterodont dentary dentition: 0, no substantial hetero- sent; 1, well-developed spine-like.
donty is present in dentary dentition; 1, single, en- 152. Scapula, blade-shape: 0, strongly expanded distally;
larged, caniform anterior dentary tooth, crown is not 1, weakly expanded, near parallel-sided.
mesiodistally expanded above root; 2, anterior dent- 153. Humeral length: 0, more than 60% of femoral length;
ary teeth are strongly recurved and caniform, but have 1, less than 60% of femoral length.
crowns expanded mesiodistally above their roots and 154. Deltopectoral crest development: 0, well-developed,
are not enlarged relative to other dentary teeth. projects anteriorly as a distinct flange; 1, rudiment-
125. Peg-like tooth located anteriorly within dentary, lacks ary, is at most a thickening on the anterolateral margin
denticles, strongly reduced in size: 0, absent; 1, present. of the humerus.
126. Alveolar foramina (‘special foramina’) medial to max- 155. Humeral shaft form, in anterior or posterior view:
illary/dentary tooth rows: 0, present; 1, absent. 0, relatively straight; 1, strongly bowed laterally along
127. Recurvature in maxillary and dentary teeth: 0, present; length.
1, absent. 156. Manual length (measured along digit ii or iii, whichever
128. Overlap of adjacent crowns in maxillary and dentary is longest) as a percentage of the combined length of
teeth: 0, absent; 1, present. the humerus and radius: 0, less than 40%; 1, more than
129. Crown is mesiodistally expanded above root in cheek 40%.
teeth: 0, absent; 1, present. 157. Metacarpals with block-like proximal ends: 0, absent;
130. Position of maximum apicobasal crown height in dent- 1, present.
ary/maxillary tooth rows: 0, anterior portion of tooth 158. Metacarpals 1 and 5: 0, substantially shorter in length
row; 1, central portion of tooth rows. than metacarpal 3; 1, subequal in length to metacarpal
131. Close-packing and quicker replacement eliminates 3.
spaces between alveolar border and crowns of adja- 159. Penultimate phalanx of the second and third fingers:
cent functional teeth: 0, absent; 1, present. 0, shorter than first phalanx; 1, longer than the first
132. Fusion between the intercentum of the atlas and the phalanx.
neural arches: 0, absent; 1, present. 160. Manual digit 3, number of phalanges: 0, four; 1, three
133. Epipophyses on anterior (postaxial) cervicals: or fewer.
0, present; 1, absent. 161. Manual digits 2–4: 0, first phalanx relatively short com-
134. Cervicals 4–9, form of central surfaces: 0, amphi- pared to second phalanx; 1, first phalanx more than
coelous; 1, at least slightly opisthocoelous. twice the length of the second phalanx.
135. Cervical number: 0, seven/eight; 1, nine; 2. ten or more. 162. Extensor pits on the dorsal surface of the distal end of
136. Articulation between the zygapophyses of dorsal ver- metacarpals and manual phalanges: 0, absent or poorly
tebrae: 0, flat; 1, tongue-and-groove. developed; 1, deep, well-developed.
137. Dorsals, number: 0, 12–13; 1, 15; 2. 16 or more. 163. Manual unguals strongly recurved with prominent
138. Sacrals, number: 0, two; 1, three; 2. four/five; 3. six or flexor tubercle: 0, absent; 1, present.
more. 164. Acetabulum: 0, at least a small perforation; 1, com-
139. Sacrum, accessory articulation with pubis: 0, absent; pletely closed.
1, present. 165. Preacetabular process, shape/length: 0, short, tab-
140. Posterior sacral ribs are considerably longer than an- shaped, distal end is posterior to pubic peduncle;
terior sacral ribs: 0, absent; 1, present. 1, elongate, strap-shaped, distal end is anterior to pubic
141. Anterior caudal vertebrae, length of transverse pro- peduncle.
cesses relative to neural spine height: 0, subequal; 166. Preacetabular process, length: 0, less than 50% of the
1, longer than neural spine. length of the ilium; 1, more than 50% of the length of
142. Proximal caudal neural spines: 0, height the same or the ilium.
up to 50% taller than the centrum; 1, more than 50% 167. Preacetabular process, lateral deflection: 0, 10–20◦
taller than the centrum. from midline; 1, more than 30◦ .
143. Elongate tail (59 or more caudals): 0, absent; 1, present. 168. Dorsal margin of preacetabular process and dorsal mar-
144. Chevron shape: 0, rod-shaped, often with slight gin of ilium above acetabulum: 0, narrow, not trans-
distal expansion; 1, strongly asymmetrically expanded versely expanded; 1, dorsal margin is transversely ex-
distally, width greater than length in mid caudals. panded to form a narrow shelf.
145. Sternal segments of the anterior dorsal ribs: 0, unossi- 169. In dorsal view preacetabular process of the ilium ex-
fied; 1, ossified. pands mediolaterally towards its distal end: 0, absent;
146. Gastralia: 0, present; 1, absent. 1, present.
147. Ossified clavicles: 0, absent; 1, present. 170. Dorsal margin of the ilium in lateral view: 0, relatively
148. Sternal plates, shape: 0, absent; 1, kidney-shaped; straight or slightly convex; 1, sinuous, postacetabular
2, shafted or hatchet-shaped (rod-like posterolateral process is strongly upturned.
process, expanded anterior end). 171. Subtriangular process extending medially from the
149. Proportions of humerus and scapula: 0, scapula longer dorsal margin of the iliac blade: 0, absent;
or subequal to the humerus; 1, humerus substantially 1, present.
longer than the scapula. 172. Subtriangular process, form and position: 0, short and
150. Scapula blade, length relative to minimum width: tab-like, above acetabulum; 1, elongate and flange-like,
0, relatively short and broad, length is 5–8 times min- on postacetabular process.
173. Brevis shelf and fossa: 0, fossa faces ventrolaterally and 197. Femoral shape in medial/lateral view: 0, bowed anteri-
shelf is near vertical and visible in lateral view along orly along length; 1, straight.
entire length, creating a deep postacetabular portion; 198. Femoral head: 0, confluent with greater trochanter,
1, fossa faces ventrally and posterior of shelf portion fossa trochanteris is groove-like; 1, fossa trochanteris
cannot be seen in lateral view. is modified into distinct constriction separating head
174. Length of the postacetabular process as a percentage of and greater trochanter.
the total length of the ilium: 0, 20% or less; 1, 25–35%; 199. ‘Anterior’ or ‘lesser’ trochanter, morphology: 0, absent;
2, more than 35%. 1, trochanteric shelf ending in a small, pointed, spike;
175. Medioventral acetabular flange of ilium, partially 2, broadened, prominent, ‘wing’ or ‘blade’ shaped, sub-
closes the acetabulum: 0, present; 1, absent. equal in anteroposterior width to greater trochanter;
176. Supra-acetabular ‘crest’ or ‘flange’: 0, present; 1, ab- 3, reduced anteroposterior width, closely appressed to
sent. the expanded greater trochanter.
177. Ischial peduncle of the ilium: 0, projects ventrally; 200. Level of most proximal point of anterior trochanter re-
1, broadly swollen, projects ventrolaterally. lative to level of proximal femoral head: 0, anterior
178. Pubic peduncle of ilium: 0, large, elongate, robust; trochanter is positioned distally on the shaft, and sep-
1, reduced in size, shorter in length than ischial ped- arated from ‘dorsolateral’ trochanter/greater trochanter
uncle. by deep notch visible in medial view; 1, anterior
179. Pubic process of ischium, shape: 0, transversely com- trochanter positioned proximally, approaches level of
pressed; 1, dorsoventrally compressed. proximal surface of femoral head, closely appressed
180. Ischium, shape of shaft: 0, relatively straight; 1, gently to ‘dorsolateral’/greater trochanter (no notch visible in
curved along length. medial view).
181. Ischial shaft, cross-section: 0, compressed mediolater- 201. Fourth trochanter of femur, shape: 0, low eminence, or
ally; 1, subcircular and bar-like. absent; 1, prominent ridge; 2, pendent.
182. Ischial shaft: 0, expands weakly, or is parallel-sided, 202. Fourth trochanter, position: 0, located entirely on prox-
distally; 1, distally expanded into a distinct ‘foot’; imal half of femur; 1, positioned at midlength, or distal
2, tapers distally. to midlength.
183. Groove on the dorsal margin of the ischium: 0, absent; 203. Anterior (extensor) intercondylar groove on distal end
1, present. of femur: 0, absent; 1, present.
184. Tab-shaped obturator process on ischium: 0, absent; 204. Posterior (flexor) intercondylar groove of the femur:
1, present. 0, fully open; 1, medial condyle inflated laterally, par-
185. Ischial symphysis, length: 0, ischium forms a median tially covers opening of flexor groove.
symphysis with the opposing blade along at least 50% 205. Lateral condyle of distal femur, position and size in
of its length; 1, ischial symphysis present distally only. ventral view: 0, positioned relatively laterally, and
186. Pubis, orientation: 0, anteroventral; 1, rotated pos- slightly narrower in width than the medial condyle;
teroventrally to lie alongside the ischium (opistho- 1, strongly inset medially, reduced in width relative to
pubic). medial condyle.
187. Shaft of pubis (postpubis), shape in cross-section: 206. Distal tibia: 0, subquadrate, posterolateral process is
0, blade-shaped; 1, rod-shaped. not substantially developed; 1, elongate posterolateral
188. Shaft of pubis (postpubis), length: 0, approximately processs, backs fibula.
equal in length to the ischium; 1, reduced, extends for 207. Fibular facet on the lateral margin of the proximal sur-
half or less the length of the ischium. face of the astragalus: 0, large; 1, reduced to small
189. Reduction of postpubic shaft: 0, postpubic shaft ex- articulation.
tends for around half the length of ischium; 1, postpubic 208. Calcaneum, proximal surface: 0, facet for tibia absent;
shaft is very short or absent. 1, well-developed facet for tibia present.
190. Body of pubis, size: 0, relatively large, makes sub- 209. Medial distal tarsal: 0, articulates distally with meta-
stantial contribution to the margin of the acetabulum; tarsal 3 only; 1, articulates distally with metatarsals 2
1, reduced in size, rudimentary, nearly excluded from and 3.
the acetabulum. 210. Metatarsal arrangement: 0, compact, closely appressed
191. Body of the pubis, massive and dorsolaterally rotated to one another along 50–70% of their length, spread
so that obturator foramen is obscured in lateral view: distally; 1, contact each other only at proximal ends,
0, absent; 1, present. spread strongly outwards distally.
192. Prepubic process: 0, absent; 1, present. 211. Digit 1: 0, metatarsal 1 robust and well-developed,
193. Prepubic process: 0, compressed mediolaterally, dor- distal end of phalanx 1–1 projects beyond the distal
soventral height exceeds mediolateral width; 1, end of metatarsal 2; 1, metatarsal 1 reduced and prox-
rod-like, mediolateral width exceeds dorsoventral imally splint-like, end of phalanx 1–1 does not extend
height. beyond the end of metatarsal 2; 2, metatarsal 1 reduced
194. Prepubic process, length: 0, stub-like and poorly de- to a vestigal splint or absent, does not bear digits.
veloped, extends only a short distance anterior to the 212. Pedal digit 4 phalangeal number: 0, five; 1, four or
pubic peduncle of the ilium; 1, elongated into distinct fewer.
anterior process. 213. Metatarsal 5, length: 0, more than 50% of metatarsal
195. Prepubic process, extends beyond distal end of 3; 1, less than 25% of metatarsal 3.
preacetabular process of ilium: 0, absent; 1, present. 214. Metatarsal 5: 0, bears digits; 1, lacks digits.
196. Extent of pubic symphysis: 0, elongate; 1, restricted to 215. Pedal unguals, shape: 0, tapering, narrow pointed,
distal end of pubic blade, or absent. claw-like; 1, wide, blunt, hoof-like.
216. Epaxial ossified tendons present along vertebral 219. Parasagittal row of dermal osteoderms on the dorsum
column: 0, absent; 1, present. of the body: 0, absent; 1, present.
217. Ossified hypaxial tendons, present on caudal vertebrae: 220. Lateral row of keeled dermal osteoderms on the dorsum
0, absent; 1, present. of the body: 0, absent; 1, present.
218. Ossified tendons, arrangement: 0, longitudinally ar- 221. U-shaped cervical/pectoral collars composed of
ranged; 1, basket-like arrangement of fusiform tendons contiguous keeled osteoderms: 0, absent; 1,
in caudal region; 2, double-layered lattice. present.
Appendix 3: Data matrix

(a = 0/1; b = 1/2; c = 2/3; d = 3/4; e = 0/2)
10 20 30 40 50 60
/ / / / / /
Euparkeria 000- - 00000 0000- 00000 000000000- - - - 000- 000 0000000000 0000000000
Marasuchus ? 0? ? ? ? ? ? ? ? ?????????? ?????????? ?????????? ?????????? ??????????
Herrerasaurus 000- - 00000 0000- 1 1 01 0 01 0000000- - - - 000- 000 000001 0000 0000000000
Abrictosaurus 000- - 1 001 1 ? ? 01 1 000? 1 00? ? 01 001 0 0000? 0- ? ? ? ?????????? ? ? ? ? 0? ? ? ? ?
Agilisaurus 000- - 1 0001 1 000- 0001 1 000? 01 0? 1 0 1 1 1 000- 000 0000000000 0? 1 0001 000
Anabisetia ?????????? ?????????? ?????1???? ?????????? ?????????? ??????????
Ankylopollexia 000- - 1 1 1 1 0 1 00- - 001 01 01 001 1 001 0 000000- 000 0000001 000 01 1 1 00000a
Ankylosauria 000- - 1 0001 1 000- 00001 1 - 0001 001 1 - 2- - 1 0- 000 01 1 0000000 0? 1 ? 1 001 00
Archaeoceratops 01 1 1 1 1 0000 ? 001 01 0001 000? ? 1 001 0 1 00021 1 1 1 0 1 1 01 ? 00001 0? 1 01 1 ? 01 0
BMNH A100 ? ? 0- - 1 ? 0? ? 1 ? 01 1 00? ? 1 00? 001 00? ? ? ? ? ? ? 0- 0? ? 00? 00? ? ? ? ? ??????????
Bugenasaura ? ? 0- - 0? 0? ? 1 ? ? ? ? 00001 01 0001 001 0 0001 00- 000 0000000? 00 01 1 000? 000
Chaoyangsaurus 1 ? 1 001 ? 00? 0? 00- 1 1 0? 1 ? ? ? ? 01 1 ? ? ? ? ? ? ? 21 0000 1 ? ? ? ? 000? ? ? ? 1 000? 0? 0
Coronosauria + 01 1 1 1 1 0000 0001 01 0001 01 0001 001 0 1 00021 1 1 1 0 01 01 1 00001 01 1 01 1 001 0
Leptoceratopsidae
Dryosauridae 000- - 1 1 1 1 0 1 1 0- - 001 01 01 001 1 01 1 0 001 000- 000 0000001 000 01 1 1 000001
Echinodon ? ? 0- - ? ? 0? ? ? ? ? ? ? 0? ? ? ? ? ? ? ? 01 ? ? ? ? ?????????? ?????????? ??????????
Emausaurus 0? 0- - ? 000? 1 000- 000? 1 000001 001 0 1 0001 0- 000 000001 0000 001 000? 000
Gasparinisaura 00? ? ? ? ? ? ? ? ? ? ? 1 0? ? ? ? 1 01 0? ? 1 001 0 0001 00- 000 0000000000 ? 01 1 00? 000
Goyocephale ? 00- - ? 001 ? 1 001 1 00001 ? ? ? ? 01 001 1 - 1 - ? 00- ? 00 1 1 1 000000- 1 ? 1 ? 00? ? ? ?
Heterodontosaurus 000- - 1 1 01 1 1 1 01 1 0001 1 001 001 001 0 000000- 001 001 00- 0000 001 0000000
Hexinlusaurus ? 00- - ? 0? ? ? ? ? ? ? ? ? ? 01 1 000? ? 1 0? 1 0 0001 00- 000 0000000000 0? 1 000? 000
Homalocephale ? 00- - ? ? ? ? ? ?????????1 1 - - ? ? 1 001 1 - 1 - - 00- 000 1 1 1 000000- 1 1 1 0000000
Hypsilophodon 000- - 1 001 1 1 01 1 00001 1 001 001 001 0 0001 00- 000 0000000000 01 1 0000001
Jeholosaurus 000- - ? ? 00? ? 01 0- 0001 1 000? 01 001 0 ? 0? 1 00- 000 0000000000 01 1 0000001
Lesothosaurus 000- - 1 0001 1 000- 000? 1 000001 01 1 0 000000- 000 000001 0000 001 0000000
Liaoceratops 0? 1 01 1 1 000 0001 01 0001 000? 01 1 0? ? ? ? ? 021 1 1 00 01 01 1 00001 01 1 000001 0
Lycorhinus ?????????? ? ? 01 1 ? ? ? ? 1 00? ? 01 0? ? ? ?????????? ?????????? ??????????
Micropachycephalosaurus ?????????? ?????????? ?????????? ?????????? ?????????? ? ? ? ? ? ? ? ? 0?
Orodromeus 000- - 1 0001 1 ? 1 0- 0001 1 01 01 01 001 0 000000- 001 000000001 0 01 1 0000000
Othnielia ?????????? ?????????? ?????????? ?????????? ?????????? ??????????
Pachycephalosauridae 000- - 1 001 1 1 001 a00001 1 - - ? 01 001 1 - 1 - - 00- 000 1 1 1 000000- 1 1 1 0000000
Parksosaurus 00? ? ? ? 0? ? ? ? ? ? 1 0? ? ? 01 01 0? ? 1 001 0 ? 0? 1 00- 000 0001 ? 00? 00 0? 1 1 00? 000
Pisanosaurus ? 0? ? ? ? ? ? ? ? ?????????? ?????1???? ?????????? ?????????? ??????????
Psittacosauridae 1 01 001 1 001 000- - 1 1 001 ? ? 0001 1 01 0 1 00? 21 01 00 000001 0000 01 1 0000000
Rhabdodontidae 000- - 1 001 0 1 00- - 00001 01 0001 00? ? ? ? ? 1 00- 000 0000001 1 00 01 1 1 000000
Scelidosaurus 000- - 1 00? ? ? 0? 0- ? ? ? 01 01 0001 001 1 - 2- 01 0- 000 01 0001 0000 001 0000000
Scutellosaurus ? 00- - ? ? 0? 1 1 ? ? ? ? 00? ? ? ? ? ? ? ? 1 0? ? ? ? ? ? ? 1 0- 000 000001 00? ? ? ? ? ? 0? ? 0? ?
Stegosauria 000- - 1 0001 1 0a0- 00001 000001 001 1 - 2- 01 0- 000 0000000000 01 1 0001 1 00
Stenopelix ?????????? ?????????? ?????????? ?????????? ?????????? ??????????
Stormbergia ?????????? ?????????? ?????????? ?????????? ?????????? ??????????
Talenkauen ? 00- - 1 01 ? ? 1 ? 01 0000? 1 0? ? ? 01 00? ? ?????????? ?????????? ??????????
Tenontosaurus dossi 000- - ? 1 1 1 0 1 001 0001 01 01 001 1 001 0 0001 00- 000 00000001 00 01 1 1 000001
Tenontosaurus tilletti 000- - 1 1 1 1 0 1 00- - 001 01 01 001 1 001 0 0001 00- 000 00000001 00 01 1 1 000001
Thescelosaurus ? 0? ? ? ? ? ? ? ? ? ? ? 1 0? ? ? ? ? ? ? ? 001 ? ? 1 0 000? 00- 000 0000000? 00 0? ? ? ? ? ? ? ? ?
Wannanosaurus ? 0? ? ? ? ? ? ? ? ?????????? ????????11 - 1 - ? 00- 000 1 01 000000- 0? 1 ? 0? ? ? ? ?
Yandusaurus ?????????? ?????????1 000001 00? ? ? ? ? ? 00- 000 0000000? ? ? ? ? ? ? ? ? ? 0? ?
Zephyrosaurus ? ? 0- - 1 ? 01 1 1 0? 1 0000? 1 0? ? 1 01 0? 1 0 1 00? 00- 001 00000? 0? 1 0 0? ? 0? ? 1 00?
Appendix 3 Continued.
70 80 90 100 110 120

/ / / / / /
Euparkeria 001 00000- - 0000001 000 00000- 0000 - - - - - - 0000 0000000000 02000000- 0
Marasuchus ?????????? ?????????0 0? 0? ? ? ? ? ? ? ?????????? ?????????? ??????????
Herrerasaurus 001 00000- - 0000001 00? 000? 0- 0000 - - - - - - 0000 0000000000 02000000- 0
Abrictosaurus ?????????? ?????????? ? ? ? ? ? ? ? ? 01 1 000- 1 0000 1 1 000? 00? 0 03001 1 ? 0- 0
Agilisaurus 001 00000- - 00001 0? 000 00? ? 0- 0001 0? 00001 000 1 1 01 000000 01 1 01 1 00- 0
Anabisetia ?????????? ?????????? ?????????? ? ? ? ? ? ? ? 01 0 1????????0 ????11?110
Ankylopollexia 1 0000000- - 00001 01 1 00 00200- 0001 1 01 01 01 01 0 1 1 1 1 1 0001 0 1 - - - 21 1 1 01
Ankylosauria 001 01 - 1 0- - 1 00000? 001 00001 1 001 1 0000- 1 1 1 00 01 01 000001 0? 1 01 1 00- 0
Archaeoceratops 002001 01 1 1 000001 ? 01 ? ? 01 ? 0- ? ? 01 1 1 01 001 000 1 1 1 1 01 1 1 00 031 01 1 ? ? ? 1
BMNH A100 ? ? ? ? 000? ? ? ? ? ? 0? ? ? ? ? ? ? ? ? ? ? ? ? ? 0? ? ? ? ? ? ? 0000 ?????????0 0301 1 1 ? 0- 0
Bugenasaura - 0000000- - 0000? ? ? ? ? ? ? ? ? ? ? ? ? ? 0? ? ? ? ? ? ? ? 000 1 1 01 1 200? 0 01 1 01 1 1 1 1 0
Chaoyangsaurus 002? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0? ? 0? ? 0- 0? 01 1 1 01 ? 01 000 ? 1 01 001 1 00 041 01 1 1 0- 0
Coronosauria + 002001 01 1 1 000001 ? 01 0 00b1 0- 1 001 1 1 01 1 01 0? 0 1 1 1 1 01 1 000 a41 01 1 1 1 1 1
Leptoceratopsidae
Dryosauridae 1 ? 000000- - 00001 01 1 00 001 00- 0001 1 01 01 01 000 1 1 1 1 000000 1 - - - 21 1 1 01
Echinodon ?????????? ?????????? ?????????1 ? ? ? ? ? 1 0000 ? ? 0? ? ? ? 0? 0 0d? ? 1 1 00- 0
Emausaurus 00? 00000- - 0000? ? ? ? 00 ? 00? ? ? ? ? 1 ? 0? ? ? ? ? 1 1 00 01 0001 0000 01 1 01 1 00- 0
Gasparinisaura ? 0? ? 00? 0- - 000000000? ? ? ? ? ? ? ? ? 0? ? ? ? ? ? ? ? 000 1 1 01 1 0001 0 ????111111
Goyocephale ? ? ? 00001 00 1 1 1 1 1 0? ? ? ? ? 1 ? ? ? ? 0? 0? 0? ? ? ? ? 0000 1 1 01 0? 01 ? 0 031 1 1 1 1 aa0
Heterodontosaurus 00200000- - 0000000000 00000- 0001 1 000- 1 0000 1 1 0000001 0 0301 1 1 1 1 01
Hexinlusaurus 00? 00000- - 00001 ? ? ? ? ? ? ? ? ? ? ? ? ? 0? ? ? ? ? ? ? 1 000 1 1 0? ? ? ? 000 ? ? ? ? 1 1 01 00
Homalocephale 0000001 1 00 1 1 1 1 1 0? 000 1 1 0? 0- ? 1 0? ?????????? ? ? ? ? ? ? ? ? 0? ????11?110
Hypsilophodon 1 1 01 0000- - 00001 00000 001 00- 0001 1 1 00001 000 1 1 01 1 0001 0 01 1 01 1 1 1 a0
Jeholosaurus 1 1 01 0000- - 00001 0? ? ? 0 ? 001 0- ? ? 01 1 1 00001 000 1 1 01 000000 00? 01 1 ? 1 1 0
Lesothosaurus 00000000- - 0000000000 000? 0- 0001 0000001 000 01 0001 0000 001 01 1 00- 0
Liaoceratops 002001 01 1 1 000001 ? 01 0 001 ? 0- 0001 1 001 001 001 1 1 1 1 001 1 00 031 01 1 1 1 1 1
Lycorhinus ?????????? ?????????? ?????????? ?????????? ?????????? ? ? ? ? 1 1 ? 0- 0
Micropachycephalosaurus ? ? 2? ? ? ? ? ? ? ?????????? ?????????? ? ? ? ? ? ? ? 000 1 1 0? ? ? ? ? ? 0 ? ? ? ? 1 1 1 0- ?
Orodromeus 0001 0000- - 00001 00000 000? ? ? ? ? 01 ? ? ? 0? 01 000 1 1 01 1 00000 01 1 01 1 00- 0
Othnielia ?????????? ?????????? ?????????? ? ? ? ? ? ? ? ? 00 ? ? ? ? ? ? ? 0? 0 ? ? ? ? 1 1 1 0- 0
Pachycephalosauridae 00001 - 1 1 00 1 1 1 1 1 0? a00 1 1 01 0- a1 01 0? ? ? ? ? 0000 1 1 01 0001 00 03aa1 1 ? aa0
Parksosaurus - 00? 0000- - ? ? 00? ? ? ? ? ? ? 0? ? ? ? ? ? 0? ? ? ? ? ? ? ? 000 1 1 0? ? ? 001 0 ????111?10
Pisanosaurus ?????????? ?????????? ?????????? ? ? ? ? ? ? ? 000 ? 1 0? 000000 ? ? ? ? 1 ? ? 0- 0
Psittacosauridae 00200001 01 00000a0000 0001 0- 1 001 1 a00001 001 1 1 1 aa00000 1- - - 111111
Rhabdodontidae - 0200000- - 00001 01 000 ? 01 ? ? ? ? ? 01 1 0001 01 01 0 1 1 1 ? ? 00000 1 - - - 21 1 1 1 0
Scelidosaurus 001 00000- - 000000? 001 001 ? 1 0001 ? 0? ? ? ? ? 1 1 00 01 01 01 0001 01 1 01 1 00- 0
Scutellosaurus ? ? 1 1 00? ? ? ? ?????????? ????????1? ? ? ? ? ? ? ? 000 0? ? ? ? 1 ? 0? 0 00? 01 1 00- 0
Stegosauria 001 00000- - 0000a01 001 00001 ? 0001 0000001 1 00 1 1 00000000 a? 1 01 1 00- 0
Stenopelix ?????????? ?????????? ?????????? ?????????? ?????????? ??????????
Stormbergia ?????????? ?????????? ?????????? ?????????? ?????????? ??????????
Talenkauen ?????????? ?????????? ? ? ? ? ? ? ? ? 01 1 1 0? 1 01 000 1 1 ? ? ? ? ? 0? 0 01 ? ? ? 1 ? ? 1 0
Tenontosaurus dossi 1 1 000000- - 00001 01 1 0? ? 0? ? 0- 0? 01 1 01 0001 01 0 1 1 1 ? ? ? 0000 0? ? ? 21 1 1 00
Tenontosaurus tilletti 1 1 000000- - 00001 01 1 00 001 ? 0- 0001 1 01 0001 01 0 1 1 1 1 1 20000 1 - - - 21 1 1 00
Thescelosaurus ? ? ? 00000- - 0000? 01 1 00 001 ? ? ? ? ? 01 ? ? ? ? ? ? 1 000 1 1 01 1 200? 0 0? 1 ? 1 1 0? 1 0
Wannanosaurus ? ? ? ? 0001 00 1 1 01 1 0? ? ? ? ? ? ? ? ? ? ? ? 0? ? ? ? ? ? ? 0000 1 1 01 0000? 0 ? ? ? ? 1 1 1 0- ?
Yandusaurus ?????????? ?????????? ? ? ? ? ? ? ? ? 0? ?????????? ?????????? ????111?10
Zephyrosaurus ? ? 01 0000- - 00? 01 00000 001 ? ? ? ? ? 0? ?????????? ?????????? 01 1 01 1 1 ? 1 0
130 140 150 160 170 180

/ / / / / /
Euparkeria 0- 1 001 0000 001 0001 000 00000? 1 000 1 1 000? 00? ? ? 0? 1 000000 0- - 201 0000
Marasuchus ?????????? ? 01 0? 0? 000 00? 0? ? ? ? ? ? ? ? 000? ? ? ? ? ? ? ? 1 000001 0- - 2000000
Herrerasaurus 0- 1 001 0000 0000201 000 00000? ? ? ? 1 1 1 0001 1 01 0 01 1 0000000 0- 01 000000
Abrictosaurus ? ? 001 ? 1 1 1 1 0? ? ? ? ? ? 2? 0 ? ? ? ? ? ? ? ? 0? 1 0000? 1 0? ? 01 ? 01 00000 0- 1 1 ? 1 00? ?
Agilisaurus 0- 02001 1 1 1 001 01 01 200 000001 0? 1 0 1 01 00? ? ? ? ? ? ? ? 01 00000 0- 01 0001 00
Anabisetia ??????111? ?????????? ? ? ? ? ? ? ? ? 00 1 001 00000? ? 0001 1 0000 0- 1 1 1 1 1 1 00
Ankylopollexia 1 01 0001 1 1 1 1 01 1 b02c ? 0 01 0001 0b00 a000000001 1 0001 0000a 0- 1 1 1 1 1 1 1 0
Ankylosauria 0- 00001 1 1 1 01 1 000? b00 000001 01 00 ? 1 000? 01 01 0001 1 1 1 1 00 0- 0000001 0
Archaeoceratops ? 1 000? 1 1 1 1 1 ? ? ? ? 1 031 0 0? ? 0? 1 ? ? ? ? ?????????? ? ? ? 01 00000 0- 1 21 1 1 1 0?
BMNH A100 ? ? 01 1 1 1 1 1 1 0? ? ? ? ? ? ? ? ? ?????????? ?????????? ?????????? ??????????
Bugenasaura 1 000001 1 1 1 0? ? ? ? ? ? ? ? ? ?????????? ?????????? ?????????? ??????????
Chaoyangsaurus 0- 00001 1 1 1 0? ? 0? ? ? ? ? ? ?????????? 1????????? ?????????? ??????????
Coronosauria + 1 1 00001 1 1 1 1 - 1 02? 03? 0 0? 0001 1 1 01 1 000000000 1 0001 00001 0- 1 21 1 1 1 1 0
Leptoceratopsidae
Dryosauridae 1 01 0001 1 1 1 0? 1 1 1 01 31 0 01 0001 01 00 00000? ? ? ? ? ? ? ? 01 00001 0- 1 1 1 1 1 1 1 0
Echinodon 0- 00? 1 1 1 1 1 0? ? ? ? ? ? ? ? ? ?????????? ?????????? ?????????? ??????????
Emausaurus 0- 00001 1 1 1 0? ? ? ? ? ? ? ? ? ?????????? ? ? ? ? ? ? 01 ? ? ?????????? ??????????
Gasparinisaura ? ? 000? 1 1 1 1 00? 0? ? ? 31 0 0? ? 1 ? 1 0? ? ? 00? 00? ? ? ? ? ? ? ? 01 00001 0- 1 1 1 1 1 1 00
Goyocephale 0- 01 ? ? 1 1 1 ? 00? ? ? 1 ? 3? 1 ? ? ? ? 01 ? 2? ? ??111????? ? ? ? 01 01 1 1 0 1 01 21 1 ? ? ? ?
Heterodontosaurus 1 a1 1 1 1 1 1 01 1 ? 001 ? 03? 0 000001 0? 01 1 00001 1 01 0 01 1 01 00000 0- 1 1 1 1 0000
Hexinlusaurus 0- 0? 001 1 1 1 001 01 01 200 000001 0? 1 0 1 000000000 00001 00000 0- 1 1 01 01 00
Homalocephale ? ? 0? ? ? 1 1 1 1 0? ? ? ? 1 ? 31 1 1 ? ? 001 ? 2? ? ?????????? ? ? ? 01 01 1 1 0 1 1 1 21 1 1 1 1 1
Hypsilophodon 1 000001 1 1 1 001 01 01 31 0 00001 1 01 00 1 000000000 00001 00000 0- 1 21 1 1 1 00
Jeholosaurus 0- 000? 1 1 1 1 001 01 0? ? ? ? 0? ? 0? 1 ? ? ? ? ?????????? ?????????? ??????????
Lesothosaurus 0- 00001 1 1 1 0000? 0? 20? ? ? ? ? ? ? ? ? 00 1 01 000000? 00001 00000 0- 01 000000
Liaoceratops 1 1 00001 1 1 1 0? ? ? ? ? ? ? ? ? ?????????? ?????????? ?????????? ??????????
Lycorhinus ? ? 0? ? 1 1 1 1 1 0? ? ? ? ? ? ? ? ? ?????????? ?????????? ?????????? ??????????
Micropachycephalosaurus 0- 00001 1 1 1 0? ? ? ? ? ? ? ? ? ? ? ? 0? ? ? ? ? ? ?????????? ? ? ? 01 0000? ?????1????
Orodromeus 0- 00001 1 1 1 001 01 01 31 0 00? 0? 1 01 00 1 0? 0000000 00001 00000 0- 1 21 1 1 1 00
Othnielia 0- 0? ? ? 1 ? 1 ? 0? ? 0? 01 3? 0 00? 01 1 01 00 1 0000? ? ? ? ? ? ? ? 01 00000 0- 1 1 1 1 1 1 00
Pachycephalosauridae 0- 0b? 0a1 1 1 0? ? ? ? 1 ? 3? 1 1 ? ? 001 ? 201 1 01 1 1 ? ? ? ? ? ? ? ? 01 01 1 1 0 1 1 1 21 1 1 1 1 1
Parksosaurus 0- 00001 1 1 1 0? ? ? ? 023? 0 0? 01 1 1 ? 1 00 00000? ? ? ? ? ? ? ? 01 00000 0- ? ? ? 1 ? ? 00
Pisanosaurus 0- ? 00? 1 1 1 1 0? ? 0? ? ? ? ? ? ?????????? ?????????? ?????????? ??????????
Psittacosauridae 1 000001 1 1 1 001 0a0031 0 0? 0001 1 1 0a 1 000000000 00001 00000 0- 1 21 1 1 1 00
Rhabdodontidae 1 01 0001 1 1 1 0? 1 1 ? 0? 3? 0 01 ? 0? 1 0? ? 0 00? 00? ? ? ? ? ? ? ? 01 001 00 0- ? 1 1 1 01 1 0
Scelidosaurus 0- 00001 1 1 1 0000002200 000001 0? 00 1 0000? ? ? ? ? ? ? ? 01 001 00 0- 00000000
Scutellosaurus 0- 0? 001 1 1 1 00? 01 01 2? ? 0? 1 ? ? ? ? ? ? ? 1 ? 000? 00? ? 00001 00000 0- ? 1 ? 0? ? 00
Stegosauria 0- 00001 1 1 1 01 1 0e0e200 01 0001 0? 00 1 1 000? 01 01 00001 1 1 1 00 0- 00000000
Stenopelix ?????????? ? ? ? ? ? ? ? 3? ? 0? ? 0? 1 ? ? ? 1 ? 0? ? ? ? ? ? ? ? ? ? ? 01 001 00 0- 1 1 1 1 01 1 0
Stormbergia ?????????? ? 0? 0? 01 200 00? ? ? ? ? ? 00 1 0? ? 0? ? ? ? ? ? ? ? 01 00000 0- 01 000000
Talenkauen ? ? ? 0? ? 1 1 1 1 0? 001 02? ? ? ? ? ? ? ? 1 0? 00 1 001 0? ? ? ? ? ? ? ? 01 00001 0- 1 1 1 1 ? 1 ? ?
Tenontosaurus dossi 1 01 0001 1 1 1 0? 01 20? 3? 0 01 1 0? 1 01 00 00000000? ? ? 0001 00001 0- 1 1 1 1 1 1 00
Tenontosaurus tilletti 1 01 0001 1 1 1 0001 20231 0 01 1 001 01 00 0000000001 1 0001 00001 0- 1 1 1 1 1 1 00
Thescelosaurus 1 000001 1 1 ? ? ? ? 0? 0231 0 01 ? 01 1 0? 00 1 000000000 00001 00001 0- 1 1 1 1 01 00
Wannanosaurus 0- 01 0? 1 1 1 1 0? ? 0? ? ? ? ? ? ?????????? ??111????? ?????????? ??????????
Yandusaurus ? ? 0? ? ? 1 1 1 1 0? ? 0? 0? ? ? ? ? ? ? ? ? ? ? ? 00 1 0? 00? ? ? ? ? ?????????? ??????????
Zephyrosaurus 0- 00? 01 1 1 1 0? ? ? ? ? ? ? ? ? ?????????? ?????????? ?????????? ??????????
190 200 210 220

/ / / /
Euparkeria 00000000- 0 00- - - 0000- 0000000000 0000000- 1 ? 0
Marasuchus 00000000- 0 00- - - 0001 0 1 00000001 0 1 001 000- 0? 0
Herrerasaurus 1 0000000- 0 00- - - 0001 0 1 000000000 0000000- 00 0
Abrictosaurus ?????????? ? ? ? ? ? ? 0? 2? 200? ? 1 ? ? ? 0 1 01 1 0? ? ? 00 0
Agilisaurus 001 1 01 1 0- 0 01 01 01 0020 200001 1 1 00 1 01 1 01 0000 0
Anabisetia 01 01 1 1 1 0- 0 01 01 1 1 01 31 201 01 1 1 1 ? 0 1 ? 1 1 01 ? 000 0
Ankylopollexia 1 1 01 1 1 1 1 00 01 01 1 1 1 1 c 1 21 1 1 1 1 1 1 00 001 1 01 0200 0
Ankylosauria 0e001 1 1 1 a1 1 1 0001 1 0- - 01 0001 1 1 - 1 21 1 1 1 1 001 1 1
Archaeoceratops 00001 1 1 1 1 0 01 1 1 01 ? 1 31 ?????11110 001 1 01 0000 0
BMNH A100 ?????????? ?????????? ?????????? ?????????? ?
Bugenasaura ?????????? ?????????? ?????????? ?????????? ?
Chaoyangsaurus ?????????? ?????????? ?????????? ?????????? ?
Coronosauria + 00001 1 1 1 1 0 01 0001 01 31 201 001 1 1 1 0 001 1 a1 0000 0
Leptoceratopsidae
Dryosauridae 1 1 01 1 1 1 0- 0 01 01 1 1 01 31 201 1 1 1 1 1 1 0 201 1 01 0000 0
Echinodon ?????????? ?????????? ?????????? ?????????? ?
Emausaurus ?????????? ?????????? ?????????? ? ? ? ? 0? ? ? 1 1 ?
Gasparinisaura 0001 1 1 1 0- 0 01 1 1 1 1 01 31 200001 1 1 1 0 201 1 01 1 000 0
Goyocephale ?????????? ?????????? ?????1??1? ? ? ? ? 01 1 1 00 0
Heterodontosaurus 0000? 1 1 0- 0 01 0001 0021 200001 ? ? 00 1 01 1 01 0000 0
Hexinlusaurus 0001 1 1 1 0- 0 01 1 1 1 1 0020 200001 1 1 1 0 1 01 1 01 0000 0
Homalocephale 00001 1 1 1 1 1 01 1 1 1 1 01 31 1 0? 0? 1 1 ? 1 0 ? ? ? ? ? 1 1 1 00 0
Hypsilophodon 0001 1 1 1 0- 0 01 1 1 1 1 01 31 200001 1 1 1 0 001 1 01 1 000 0
Jeholosaurus ?????????? ? ? ? ? ? ? 01 31 200001 1 1 1 0 1 01 1 0? ? ? 00 0
Lesothosaurus 001 001 1 0- 0 01 0001 0020 200001 1 1 00 1 ? 1 1 01 0000 0
Liaoceratops ?????????? ?????????? ?????????? ?????????? ?
Lycorhinus ?????????? ?????????? ?????????? ?????????? ?
Micropachycephalosaurus ?????????? ? ? ? ? ? ? 01 31 ? 0? ? ? 1 ? ? ? ? ?????????? ?
Orodromeus 0001 1 1 1 0- 0 01 1 1 1 1 01 31 2000a1 1 1 1 0 001 1 01 ? 000 0
Othnielia 0001 1 1 1 0- 0 01 1 1 1 1 01 31 200001 1 1 1 0 1 01 1 01 ? 000 0
Pachycephalosauridae 00001 1 1 1 1 1 01 1 1 1 1 01 31 1 0? 0? ? ? ? ? ? ? ? ? ? ? 1 1 1 00 0
Parksosaurus 0001 1 1 1 ? ? 0 01 1 1 1 1 01 31 21 00? 1 ? ? ? 0 001 1 01 1 000 0
Pisanosaurus ?????????0 ?????????? ? ? ? ? ? 0? 0? 0 ? ? ? ? 0? ? ? 00 0
Psittacosauridae 00001 1 1 1 1 0 01 1 1 01 01 31 201 001 1 1 00 001 1 01 0000 0
Rhabdodontidae 1 0001 ? ? ? ? ? ? ? ? ? ? ? 1 1 31 21 1 1 ? 1 1 1 ? ? ? ? ? ? ? 1 1 ? 00 0
Scelidosaurus 00001 1 1 0- 0 1 1 0001 0020 21 0001 1 1 00 001 1 1 1 001 1 1
Scutellosaurus 000001 1 0- 0 01 0? ? 1 0020 200001 ? ? ? 0 ? ? ? ? 01 ? ? 1 1 0
Stegosauria 02001 1 1 0- 0 a1 01 01 1 0- - 01 0001 1 1 - 1 21 1 1 1 a001 1 0
Stenopelix 00001 1 ? ? ? 0 01 1 1 a1 01 c 1 ?????1???0 001 1 01 0000 0
Stormbergia 00a1 1 1 1 0- 0 01 0001 0020 200001 1 1 ? 0 1 ? 1 1 01 ? ? 00 0
Talenkauen ?????11??0 01 01 1 1 ? 1 31 2? ? ? ? 1 1 1 ? 0 00? 1 01 ? 000 0
Tenontosaurus dossi 01 01 1 1 1 0- 0 01 01 1 1 ? 1 31 21 1 1 1 1 1 1 ? 0 001 1 01 1 000 0
Tenontosaurus tilletti 01 01 1 1 1 1 00 01 01 1 1 1 1 31 21 1 1 1 1 1 1 1 0 001 1 01 1 000 0
Thescelosaurus 0001 1 1 1 0- 0 01 1 1 1 1 01 31 21 0001 1 1 1 0 001 1 01 1 000 0
Wannanosaurus ?????????? ? ? ? ? ? ? 0? ? ? 1 0? 0? 1 ? ? ? ? ? ? ? ? ? ? ? ? 00 0
Yandusaurus ?????????? ?????????? ? ? 000? ? ? ? ? ? ? ? ? 0? ? ? ? ? ?
Zephyrosaurus ?????????? ?????????? ?????????? ?????????? ?
Appendix 4: Tree descriptions Node 15 (Cerapoda): Unambiguous: 174 (1 to 2), 211 (1 to

0); DELTRAN: 52, 63 (1 to 0), 92, 119, 139.
The ‘describetrees’ option of PAUP∗ was used to interpret
character state transformations. All transformations are based Node 16 (Ornithopoda): Unambiguous: 13, 64, 105;
upon the derivative strict reduced consensus tree (see Fig. 4). ACCTRAN: 217.
Transformation was evaluated under accelerated transforma- Node 17: Unambiguous: 34, 6 1, 118; ACCTRAN: 9, 60, 62,
tion (ACCTRAN) and delayed transformation (DELTRAN) 83, 121; DELTRAN: 91, 145, 217.
options; unambiguous synapomorphies are those that dia-
Node 18: Unambiguous: 19 (1 to 0), 22, 64 (1 to 0), 137 (1 to
gnose a node under both ACCTRAN and DELTRAN optim-
2), 142, 174 (2 to 1); ACCTRAN: 10 (1 to 0), 13 (1 to 0), 54,
isation. Node numbers refer to Fig. 4. For simple 0–1 state
60 (1 to 0), 62 (1 to 0), 77, 92 (1 to 0), 95, 202; DELTRAN:
changes only the character number is given; for other state
14, 83, 121.
changes the type of change is specified in parentheses.
Node 19: Unambiguous: 99, 193 (1 to 0), 203, 205;
Node 1 (Ornithischia): Unambiguous: 26, 102, 115, 127, ACCTRAN: 47, 48, 103, 111, 123, 134, 145 (1 to 0), 160,
128, 129, 130; ACCTRAN: 6, 10, 11, 20, 29, 53, 77 (1 to 0), 161, 182.
90, 101, 112 (2 to 0), 116, 123 (1 to 0), 138 (0 to 2), 152 (1 Node 20: Unambiguous: 115 (1 to 2), 151 (1 to 0), 197, 204;
to 0), 165, 186, 187, 192, 196, 199 (1 to 2), 201 (1 to 2), 207, ACCTRAN: 179, 181; DELTRAN: 9, 10 (1 to 0), 13 (1 to
211, 213, 214, 216. 0), 54, 77, 92 (1 to 0), 103, 111, 123, 134, 202.
Node 2 (Heterodontosauridae + Genasauria): Unambigu- Node 21: Unambiguous: 7, 8, 18, 25, 6 0, 78, 93, 119 (1 to
ous: 206; ACCTRAN: 208; DELTRAN: 6, 10, 11, 20, 29, 0), 170; DELTRAN: 145 (1 to 0), 160, 161, 182.
53, 77 (1 to 0), 90, 116, 123 (1 to 0), 135, 138 (0 to 2), 152
(1 to 0), 165, 186, 187, 192, 196, 199 (1 to 2), 201 (1 to 2), Node 22:Unambiguous: 62, 133 (1 to 0), 135 (1 to 2), 143;
206, 211, 213, 214, 216. ACCTRAN: 47 (1 to 0), 95 (1 to 0), 106 (0 to 2), 179 (1 to
0), 181 (1 to 0); DELTRAN: 48
Node 3 (Heterodontosauridae): Unambiguous: 9, 14, 15,
91, 96, 125, 173, 176; ACCTRAN: 12, 23, 40, 43, 6 3 (1 Node 23: Unambiguous: 34 (1 to 0), 120, 217 (1 to 0);
to 2), 109, 112 (0 to 3), 117, 121, 137 (1 to 0), 175, 200; ACCTRAN: 48 (1 to 0); DELTRAN: 47, 95, 179, 181.
DELTRAN: 101, 112 (2 to 3), 157, 162. Node 24: Unambiguous: 109, 144, 151 (1 to 0); ACCTRAN:
Node 4: Unambiguous: 114, 124; ACCTRAN: 7, 138 (2 to 52 (1 to 0), 75 (1 to 0), 77 (1 to 0), 121 (1 to 0); DELTRAN:
3). 54.
Node 5 (Genasauria): Unambiguous: 97, 113, 126 (1 to 0), Node 25 (Marginocephalia): Unambiguous: 19 (1 to 0), 68,
156 (1 to 0), 159 (1 to 0), 163 (1 to 0); ACCTRAN: 150 (1 84, 112 (1 to 3), 150, 184 (1 to 0), 188, 189; ACCTRAN: 31,
to 0), 157 (1 to 0), 162 (1 to 0); DELTRAN: 148, 207, 208. 41, 94, 108, 136, 137 (1 to 0), 145 (1 to 0), 147, 203.
Node 6 (Thyreophora): Unambiguous: 106; ACCTRAN: Node 26 (Ceratopsia): Unambiguous: 3, 11 (1 to 0), 16, 27,
19 (1 to 0), 101 (1 to 0), 160, DELTRAN: 46, 112 (2 to 0). 35 (0 to 2), 36, 63 (0 to 2); ACCTRAN: 1, 7, 17, 70, 75 (1 to
0), 76, 107, 195 (1 to 0); DELTRAN: 91.
Node 7: Unambiguous: 35, 89, 219, 220; ACCTRAN: 31,
77, 85, 211 (1 to 0). Node 27: Unambiguous: 38, 103, 118, 120, 121; ACCTRAN:
38, 41 (1 to 0), 87, 100; DELTRAN: 31, 70, 75 (1 to 0), 137
Node 8: Unambiguous: 98, 112, 158; ACCTRAN: 135 (1 to (1 to 0), 147, 195 (1 to 0), 203.
0), 137 (1 to 2), 168, 174 (1 to 0), 185, 191, 202, 221.
Node 28 (Neoceratopsia): Unambiguous: 5, 10 (1 to 0), 37,
Node 9: Unambiguous: 30, 32 (0 to 2), 80, 215; ACCTRAN: 42, 44, 45, 50, 59, 66, 69, 79, 83, 122; ACCTRAN: 1 (1 to
42, 104, 110; DELTRAN: 19 (1 to 0), 85, 135 (1 to 0), 137 0), 2, 17 (1 to 0), 135 (1 to 2), 16 1, DELTRAN: 14, 76, 94,
(1 to 2), 168, 174 (1 to 0), 185, 191, 202. 107.
Node 10 (Eurypoda): Unambiguous: 46 (1 to 0), 58, 106 (1 Node 29: Unambiguous: 4, 39, 55, 56, 106, 131; ACCTRAN:
to 0), 132, 133, 152, 166, 167, 197, 201 (2 to 0), 210, 211, 4, 7 (1 to 0), 100 (1 to 0); DELTRAN: 2, 27 (1 to 0).
212; ACCTRAN: 52, 86, 182 (0 to 2); DELTRAN: 160.
Node 30 (Pachycephalosauria): Unambiguous: 30, 32, 43,
Node 11 (Neornithischia): Unambiguous: 184; ACCTRAN: 71, 72, 74, 124, 153, 154, 155, 168, 179, 201; ACCTRAN:
46, 52, 75, 92, 104, 112, 133, 185. 9, 15, 21, 81, 82, 88, 91 (1 to 0), 114, 140, 148 (1 to 2);
Node 12: Unambiguous: 178, 194; ACCTRAN: 149; DEL- DELTRAN: 41, 97 (1 to 0).
TRAN: 75, 101, 104, 112 (2 to 1), 133. Node 31: Unambiguous: 42, 51, 73, 167, 169, 171, 217, 218;
Node 13: Unambiguous: 173, 176, 193, 195, 209; ACCTRAN: 118, 141, 180, 190; DELTRAN: 9, 14, 82, 108,
ACCTRAN: 14, 63 (1 to 0), 91; DELTRAN: 185. 136, 140, 148 (1 to 2).
Node 14: Unambiguous: 117, 138 (2 to 3), 175, 177, 198, Node 32: Unambiguous: 67, 172; DELTRAN: 21, 81, 88,
199 (2 to 3), 200; ACCTRAN: 119, 139, 145, 149 (1 to 0). 141, 180, 190.

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Journal of Systematic Palaeontology 6 (1): 1–40 Issued 22 February 2008

doi:10.1017/S1477201907002271 Printed in the United Kingdom

The phylogeny of the ornithischian

KEY WORDS vertebrate palaeontology, Ornithischia, systematics, cladistics, Dinosauria

∗ E-mail: [email protected]; †Email: [email protected]; ‡Email: [email protected]

Introduction tebrate faunas are dominated by saurischians (Weishampel

Institutional abbreviations ZDM = Zigong Dinosaur Museum, Dashanpu,

Clade name Phylogenetic deﬁnition

(Appendix 4) indicates that Marginocephalia is supported by ceratopsian node (including Micropachycephalosaurus) is

Appendix 3: Data matrix

70 80 90 100 110 120

130 140 150 160 170 180

190 200 210 220

Appendix 4: Tree descriptions Node 15 (Cerapoda): Unambiguous: 174 (1 to 2), 211 (1 to

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