Yates

Solving a dinosaurian puzzle: the identity of Aliwalia rex Galton

ADAM M. YATES

Bernard Price Institute for Palaeontological Research, University of the Witwatersrand, Private Bag 3, WITS 2050,

South Africa

AbstractEucnemesaurus fortis Van Hoepen 1920 from the Late Triassic of South Africa is demonstrated to be the senior synonym of thepuzzling dinosaur taxon Aliwalia rex Galton 1985. A new specimen of this poorly-known taxon is described. Eucnemesaurus isclearly a sauropodomorph and increases the diversity of sauropodomorph taxa in the South African Late Triassic to six. Itshares a number of femoral synapomorphies with Riojasaurus from the Late Triassic of Argentina and Riojasauridae tax. nov. iserected to accommodate them. These conclusions are supported by a comprehensive cladistic analysis of 46 sauropodomorphand other basal dinosauriform taxa using 353 osteological characters. This analysis also supports the paraphyletic nature of thetraditional ‘prosauropod’ assemblage.

Keywords: Aliwalia, Eucnemesaurus, Late Triassic, Sauropodomorpha, South Africa, Riojasauridae

Introduction

Alfred “Gogga” Brown discovered the first Triassic

dinosaurs from South Africa sometime during the

early 1860’s. His early collections were all made at

“Barnard’s Spruit, Ward, 15 miles south of Aliwal

North” (Galton and Van Heerden 1998) and were

collected from a unit that is now called the lower

Elliot Formation. The fossils were sent in five

shipments to various museums in Europe (Seeley

1894; Broom 1911). The bulk of these collections

consisted of moderately large sauropodomorph

dinosaurs, which are the commonest fossils in the

lower Elliot Formation. However the shipment that

went to the Naturhistorisches Museum in Vienna

contained a peculiar dinosaur femur that is from a

decidedly rare taxon. Indeed, until now, this femur

was thought to be unique, although it has been

suggested that a maxilla of a carnivorous archosaur,

included amongst the bones that Brown sent to

London, might also belong to the same taxon as the

femur (Galton 1985).

Von Huene was the first to describe this particular

femur and he tentatively placed it in the genus

Euskelosaurus (Huene 1906). Euskelosaurus browni

Huxley 1866 was the name given to the large

sauropodomorph bones in the first shipment that

Brown had sent to London (Huxley 1866). The name

is no longer regarded as valid since the lectotype of E.

browni displays no diagnostic characters beyond

Sauropodomorpha (Yates 2003; Yates and Kitching

2003). Cooper went further than Huene and

suggested that the Vienna femur was part of the

same individual that included the type series of

E. browni (Cooper 1980). Galton pointed out that it

was quite distinct from the lectotype femur of E.

browni and indeed appeared to be quite distinct from

all other sauropodomorphs (Galton 1985). Conse-

quently he erected Aliwalia rex Galton 1985 to

accommodate it (Galton 1985). The systematic

position of A. rex has always been uncertain. Galton

suggested that it belonged to an herrerasaurid which

was the position taken by Paul (1988). The belief that

A. rex represented a large, carnivorous dinosaur,

whether or not it was an herrerasaurid, was the part of

the basis for referring a maxilla of a large carnivorous

archosaur to this taxon (Galton 1985; Galton and Van

Heerden 1998). This referral was also supported by the

fact that the maxilla was part of the Brown’s London

ISSN 0891-2963 print/ISSN 1029-2381 online q 2006 Taylor & Francis

DOI: 10.1080/08912960600866953

Correspondence: A.M. Yates, Bernard Price Institute for Palaeontological Research, University of the Witwatersrand, Private Bag 3, WITS2050, South Africa. E-mail: [email protected]

Historical Biology, 2006; 1–31, iFirst article

shipment of bones and had therefore been found at the

same locality as the lectotype femur. Sues commented

that there were no shared derived characters that link

A. rex to Herrerasaurus and that it was best classified as

Dinosauria incertae sedis (Sues 1990), a position that

was agreed upon by later workers (Galton and Van

Heerden 1998; Langer 2004a).

In 2003 a party of geologists and palaeontologists

from the University of Witwatersrand found a small

assemblage of dinosaur bones in the lower Elliot

Formation, on the farm Spioenkop, Rosendal District,

Free State South Africa. This assemblage included the

proximal and distal end of a femur that was quickly

recognised as Aliwalia rex, the second specimen

recognised since the initial discovery over 130 years

previously. It was with anticipation of solving the

riddle of the identity of A. rex that the associated bones

were prepared. Unexpectedly the dorsal vertebra had

diagnostic synapomorphies of Sauropodomorpha and

did not differ any noticeable way from the large basal

sauropodomorphs common in the lower Elliot

Formation. Examination of the numerous scrappy

holotypes of dinosaurian taxa named from the lower

Elliot Formation over the years has revealed that this is

not the first time an ‘Aliwalia-type’ femur has been

found associated with sauropodomorph vertebrae.

Specifically the long-forgotten Eucnemesaurus fortis

Van Hoepen 1920 which had been dismissed as either

a synonym of Euskelosaurus browni or a nomen

dubium, is a third example of an ‘Aliwalia-type’

femur, once again associated with typical basal

sauropodomorph vertebrae (Van Hoepen 1920). In

this case it appears that the accumulation represents

the fragmentary remains of a pelvis, hind leg and tail of

a single individual. This paper outlines the evidence

that Aliwalia rex is a junior synonym of Eucnemesaurus

fortis and that the species is a sauropodomorph with

close affinities to Riojasaurus incertus from the Late

Triassic of Argentina.

Institutional abbreviations: BP, Bernard Price

Institute, University of the Witwatersrand, Johannes-

burg; NMW, Naturhistorisches Museum, Vienna;

PVL Instituto Miguel Lillo, Tucuman; SMNS,

Staatliches Museum fur Naturkunde, Stuttgart; TM,

Transvaal Museum, Pretoria.

The synonymy of Eucnemesaurus fortis and

Aliwalia rex

Although much of TM 119 (the holotype of

Eucnemesaurus fortis) could not be located during a

visit to the Transvaal Museum (the distal end of the

pubis, a section of the femur and the tibia were

missing), the morphology of the material is well

recorded by photographs and accurate (if minimalist)

line drawings in Van Hoepen (1920) (Figure 1). These

show clearly that, like NMW 1889-XV-39 (the

holotype femur of Aliwalia rex), the femoral head is

elongate for a basal sauropodomorph, that there is a

large proximal tubercle on the posterior surface, the

lesser trochanter is developed into a tall crest and the

long axis of the base of the fourth trochanter extends

obliquely from a position against the medial margin of

the shaft in posterior view, at its proximal end, to a

position more centrally placed on the posterior surface

of the shaft, distally. The large posterior tubercle is an

especially unusual character that represents a reversal

to the non-dinosaurian condition (Novas 1996). The

only other dinosaur to exhibit one is Riojasaurus

incertus. Points of dissimilarity between the two

holotypes are probably caused by damage to one, or

the other, of the specimens. In TM 119 the proximal

end of the lesser trochanter ends abruptly whereas the

proximal end slopes more gradually toward the shaft

in NMW 1889-XV-39. However the proximal end of

the lesser trochanter of both specimens has been

damaged and the tip of the trochanter could simply

have been lost in the latter specimen. The well-

preserved lesser trochanter of BP/1/6111 demon-

strates that the proximal end terminates abruptly. In

TM 119 and NMW 1889-XV-39 the fourth trochan-

ter is relatively proximally placed although it is more

extreme in NMW 1889-XV-39. In TM 119 the first

rise of the proximal end of the fourth trochanter was

placed 175 mm from the proximal end of the bone

according to Van Hoepen (1920). This distance is

1.13 times the length of the proximal head of the bone,

whereas in NMW 1889-XV-39 this distance is only

0.95 of the length of the proximal head. This

difference may be simple individual variation but

there are reasons to suspect that the fragment of

NMW 1889-XV-39 has not been correctly attached to

the proximal end of the bone. There is a large obvious

repaired join between the two pieces and the medial

margin of the piece bearing the fourth trochanter is

strongly deflected medially relative to the long axis of

the femoral head. Since the medial margin of the

femur in the vicinity of the fourth trochanter is more

or less vertical in other dinosaurian femora it would

imply that the head of NMW 1889-XV-39 would be

angled strongly downwards. Such an arrangement is

not impossible but seems unlikely in a moderately

derived sauropodomorph. It is more likely that the two

pieces are mismatched and the head of the femur was

angled medially as in other sauropodomorphs

(Figure 2). The base of the fourth trochanter in TM

119 is figured (in posterior view) as starting at the

medial edge at its proximal end and curving in away

from it at its distal end (Van Hoepen 1920). This is

less apparent in NMW 1889-XV-39 but it is still

somewhat obliquely oriented and, as explained above,

the fragment that bears the fourth trochanter in this

specimen is probably displaced. The new femur,

BP/1/6111, clearly displays a medio-distally curving

fourth trochanter as well as a rounded profile of the

A. M. Yates2

Figure 2. Proximal ends of the femora of Plateosaurus engelhardti and Eucnemesaurus fortis in posterior view. A, Plateosaurus engelhardti, SMNS

13200. B, Eucnemesaurus fortis, TM 119, redrawn from Van Hoepen (1920). C, Eucnemesaurus fortis, NMW 1889-XV-39, as restored.

D, Eucnemesaurus fortis, NMW 1889-XV-39, with the position of the distal fragment shifted to bring the specimen into line with A and

B. Each figure displays the angle between the medial margin of the shaft and the axis running from the proximolateral corner of the femur to

the peak of the femoral head. Numbers below each figure indicate the distance between the proximal end of the bone and the proximal end of

the fourth trochanter divided by the proximal length of the femoral head. Grey areas represent restored areas, black areas represent broken

bone surfaces. Scale bars represent 100 mm.

Figure 1. Unlocated bones from the holotype of Eucnemesaurus fortis as figured by Van Hoepen (1920). Distal fragment of the left pubis in

proximal view showing a section through the pubic blade (A) and distal view (B). Proximal end of the left femur in lateral (C), and posterior

(D) views. Left tibia in lateral (E), posterior (F), proximal (G) and distal views (H). Scale bar represents 100 mm.

Solving a dinosaurian puzzle 3

fourth trochanter and a notched (‘semi-pendant’)

distal end. The latter two characters are present in

NMW 1889-XV-39 but cannot be determined in TM

119 because the fourth trochanter is broken off at

its base.

The occurrence of this constellation of unusual

characters (where they can be determined) in all three

specimens from the same stratigraphic unit is good

evidence that they belong to a single taxon. The only

major difference between NMW 1889-XV-39 and

TM 119 is the relative position of the fourth

trochanter but this may well be an artefact of

restoration in the holotype of Aliwalia.

Systematic palaeontology

Sauropodomorpha Huene, 1932

Massopoda taxon nov.

Definition: The most inclusive clade containing Salt-

asaurus loricatus but not Plateosaurus engelhardti. As it

currently stands this taxon is a homodefinitional

synonym of Sauropoda but as explained below the

content of Sauropoda in the phylogeny supported here

(and elsewhere) is radically expanded beyond all

traditional usage of the name Sauropoda and a

redefinition of this taxon is required.

Etymology: Massa (Latin), lump and pous (Greek),

foot. Also a contraction of Massospondylidae and

Sauropoda, two well-known and disparate subordi-

nate taxa within Massopoda.

Riojasauridae taxon nov.

Definition: The most inclusive clade containing

Riojasaurus incertus but not Plateosaurus engelhardti,

Massospondylus carinatus, or Anchisaurus polyzelus. As

undesirable as multiple anchor taxa may be, it is

necessary to use them here to preserve the intent of the

name across the widely differing hypotheses of

sauropodomorph phylogeny that have been proposed

recently. It is important to note that Melanorosaurus

readi was deliberately left out of the list of exclusive

anchors. This is because it makes little sense to

differentiate Riojasauridae from Melanorosauridae in

situations where these taxa form a small low-diversity

clade within Sauropodomorpha such as in Galton and

Upchurch (2004). In cases like these Riojasauridae

simply becomes a heterodefinitional junior synonym

of Melanorosauridae as defined by Galton and

Upchurch (2004). However, in the phylogenetic

hypothesis favoured here, Riojasauridae and Mela-

norosauridae are distinct and distantly related taxa

amongst basal sauropodomorphs.

Included Taxa: Riojasaurus incertus and Eucnemesaurus

fortis

Diagnosis: Large, robust sauropodomorphs with

the following synapomorphies: a tall, crest-like lesser

trochanter that is higher than wide in cross-section;

a large tubercle on the posterior side of the proximal

femur (this is a reversal to a non-dinosaurian

character-state); and a rounded, convex profile of

the fourth trochanter (versus a subrectangular

profile). The presence of a shallow embayment on

the distal margin of the fourth trochanter (giving the

trochanter a semi-pendant shape) might also diagnose

this clade but the presence of this character in

Massospondylidae renders its distribution ambiguous.

Eucnemesaurus fortis Van Hoepen 1920

1906 ? Euskelosaurus Huene (1906), pp. 131–132,

Figures 41–42, pl. 16, Figure 2, pl. 17,

Figure 1

1920 Eucnemesaurus fortis Van Hoepen (1920),

pp. 93–102, Figures 1–7, pl. 11–13

1979 cf. Euskelosaurus Van Heerden (1979), p. 69,

Figures 19–20, pl. 52–55

1985 Aliwalia rex Galton (1985), pp. 15–16,

Figure 5d–i, pl. 5

Diagnosis: A sauropodomorph with: a small accessory

lamina branching off of the paradiapophyseal lamina

and dividing the middle chonos in the middle dorsal

vertebrae; an abrupt proximal end of the lesser

trochanter; a fourth trochanter with a curved and

oblique long axis .

Holotype: TM 119 a fragmentary posterior postcranial

skeleton including two incomplete dorsal neural

arches, a dorsal centrum, distal and proximal

fragments of the left pubis, the proximal end of the

left femur, the left tibia and four incomplete proximal

to mid caudal vertebrae. From Farm Zonderhout,

near Slabberts, Free State, South Africa, lower Elliot

Formation (Norian, Late Triassic).

Referred Material: NMW 1889-XV-39 and 1876-VII-

B124. Proximal and distal ends of a left femur.

Holotype of Aliwalia rex. From Barnard’s Spruit,

Aliwal North District, Eastern Cape, South Africa,

lower Elliot Formation.

BP/1/6107, 6110–6115, 6220. Incomplete post-

cranial skeleton including a posterior dorsal vertebra

(6107), left coracoid (6113), a ventral fragment of a

left scapula (6114), a fragment of the dorsal end of

a left scapula? (6115), a proximal caudal vertebra

(6220), fragments of a right proximal femur (6111),

distal end of a right femur (6110) and two rib shaft

fragments (6112). The bones have been catalogued

individually because the remains were disarticulated

(although in close proximity to one another) and

the association cannot be proven. Nevertheless it

A. M. Yates4

seems likely that these bones come from a single

skeleton because of the lack of duplication, and the

similarity in size, and state of preservation. In

addition several weathered pieces were found

downstream from the site (BP/1/6151). These

may, or may not, belong to the specimen. They

include fragmentary mid and distal caudal vertebrae

and two phalanx fragments. From the dongas

(erosional gullies) at the base of Spioenkop, Heelbo

farms, Rosendal District, Free State South Africa.

The specimens were collected in dark purple-grey

silts in the lower Elliot Formation.

Description

Van Hoepen (1920) described the holotype specimen

so this description concentrates on the new material in

the BP collections. Nevertheless certain aspects of the

holotype are described here in light of recent advances

in our knowledge of characters significant to sauro-

podomorph phylogeny. The Vienna specimen has

been described elsewhere (Galton 1985; Galton and

Van Heerden 1998).

Dorsal vertebrae (Figure 3). There are two incomplete

middle to posterior dorsal neural arches of TM 119

that are available for study, with the middle dorsal

being the more informative of the two. It has a laterally

directed transverse process on the right side from

which arise three diapophyseal lamiae: the

postzygodiapophyseal, posterior centrodiapophyseal

and paradiapophyseal laminae. These laminae bound

the posterior and middle chonae. A unique accessory

lamina branches off of the paradiapophyseal lamina

and extends ventrally for a short distance. It does not

reach the neurocentral suture but does divide the

middle chonos into two compartments. As in all other

early sauropodomorphs the prezygodiapophyseal

lamina and its associated anterior chonos are

absent, although there is a weak horizontal ridge

extending from the diapophysis to the base of the

prezygapophysis. The zygopophyses have simple

rounded margins in dorsal view and are horizontally

oriented. The posterior dorsal neural arch preserves

the base of a shallow, triangular hyposphene

that extends ventrally from between the two

postzygapophyses. The position of the ventral margin

Figure 3. Dorsal vertebrae of Eucnemesaurus fortis. A, Middle dorsal neural arch (TM 119) in left lateral view. B, Posterior dorsal neural arch

(TM 119) in posterior view. Posterior dorsal vertebra (BP/1/6107) in left lateral (C), anterior (D) and posterior (E) views. Scale bars represent

50 mm. Abbreviations: a.l.mc, accessory lamina of middle chonos; a.l.pc, accessory lamina of the post chonos; c, centrum; dp, diapophysis; hs,

hyposphene; n.c, neural canal; n.sp, neural spine; pcd.l posterior centrodiapophyseal lamina; pp, parapophysis; pod.l, postzygapophyseal

lamina; poz, postzygapophyses; ppd.l, paradiapophyseal lamina; prz, prezygapophysis; t.p, transverse process.


Figure 5. Left pectoral girdle of Eucnemesaurus fortis. Ventral end of the scapula (BP/1/6114) and complete coracoid (BP/1/6113) in lateral

(A), medial (B) and ventral (C) views. D, Dorsal fragment of left scapula (BP/1/6115) in lateral view. Scale bar represents 50 mm.

Abbreviations: a.f, acromial fossa; c.f, coracoid foramen; c.t, coracoid tubercle; g, glenoid; s.b, scapula blade; s.m.t, scar for scapula head of

m. triceps.

Figure 4. Caudal vertebrae of Eucnemesaurus fortis. Proximal caudal vertebra (BP/1/6220) in anterior (A), posterior (B), left lateral (C),

dorsal (D) and ventral (E) views. F, Middle caudal vertebra (TM 119) in right lateral view. Scale bars represent 50 mm. Abbreviations: a.t,

anterior tubercle; c.f, chevron facet; p.r, parasagittal ridge; s.r sagittal ridge, other abbreviations as in Figure 2.

A. M. Yates6

indicates that the size of the articular surfaces of the

hyposphene would have been much smaller than

the postzygapophyseal facets. Only the lower part

of the neural spine is preserved but it is enough to show

that the spine was a simple, transversely compressed

laminar structure.

There is a posterior dorsal vertebra in the

Spioenkop assemblage. It probably represents the

thirteenth or fourteenth dorsal based on the position

of the parapophysis. Unfortunately the prezygapo-

physis, postzygapophyses, diapophyses, neural spine

and hyposphene have all broken off at their base. The

centrum is massive, amphicoelous and lacks pleur-

ocoels, or even pleural fossae, on each side. It is 98 mm

long, which is slightly less than its maximum height of

106 mm. The anterior and posterior centrum faces are

also slightly higher than they are wide. The neural arch

is low with the height from the postzygapophyseal

facets to the neurocentral sutures is much less than the

height of the centrum. The middle and posterior

chonae are impressed below the base transverse

process and are separated by a well-developed

posterior centrodiapophyseal lamina. The paradia-

pophyseal lamina is not present because the parapo-

physis abuts the base of the transverse process. There

is a smooth surface between the parapophysis, the

base of the transverse process and the base of the

prezygapophysis, which lacks both an anterior chonos

and the prezygodiapophyseal lamina. A well-devel-

oped hyposphene is present below the postzygapo-

physeal facets and is 75% of the height of the

subcircular neural canal. The base of the neural spine

is anteroposteriorly elongate and transversely

compressed.

Caudal vertebrae (Figure 4). A proximal caudal

vertebra is present in the Spioenkop collection, while

there are two proximal caudals, a middle caudal and a

distal caudal present in TM 119. The Spioenkop

proximal caudal (BP/1/6220) is well preserved,

although it is missing the right transverse process,

the dorsal tip of the neural spine and the tips of both

prezygapophyses. The gently amphicoelous centrum

is very slightly wedge shaped in lateral view, with a

length of 75 mm at the ventral end and 80 mm

dorsally. It is also taller than it is long with a maximum

height of 90 mm. The mid-length cross-section is

ovoid and dorso-ventrally tall, with a transversely

rounded ventral surface. The central faces are also

Figure 6. Pelvic and hindlimb bones of the holotype of Eucnemesaurus fortis (TM 119). Proximal end of left pubis in lateral (A) and anterior

(B) views. Proximal end of left femur in lateral (C) anterior (D) and proximal (E) view. Scale bars represent 50 mm. Abbreviations: f.h, femoral

head; i.a.s, ilial articular surface; l.t, lesser trochanter; o.f, obturator fenestra; p.b, pubic blade; p.t, posterior tubercle.


dorsoventrally taller than they are wide with the

anterior face being 82 mm high and 67 mm wide and

the posterior face being 90 mm high and 68 mm wide.

The ventral portion of this (16 mm deep at the

midline) forms a convex chevron facet that is oriented

posteroventrally. The transverse processes have flatly

elliptical bases that are oriented horizontally. The

flattened process is backswept and slightly dorsally

directed. The neural spine is a simple, transversely

flattened rod. Its anteroposterior length (measured

above the postzygapophyseal buttresses is more than

half the length of the neural arch at its junction with

the centrum. A midline keel begins at the base of the

anterior margin of the neural spine and extends

forwards to the interprezygapophyseal space, where

it terminates in a small raised tubercle. There are

shallow paramedian fossae on each side of this

keel that are flanked by short, low ridges. The

postzygapophyses project posterior to the neural spine

and there is a small notch developed between them.

The ventral margins of the postzygapophyseal facets

are separated and there is no hyposphenal ridge.

The proximal caudal from TM 119 is less

completely preserved and differs only in that the

anterior central face is not as strongly laterally

compressed as in BP/1/6220. In the former specimen

the width of the anterior centrum face is 94% of its

height while it is 82% in the former. This is probably

due simply to individual variation. The other proximal

caudal is from a more distal position in the tail and has

a centrum that is longer than high (80 mm long and

65 mm high) The ventral surface of the centrum is

broad and rounded and lacks any trace of a midline

sulcus. The anterior centrum face is as wide as it is

high. The anterior median ridge is poorly developed

and there are no paramedian fossae or ridges. The

more distal caudals do not record any significantly

different features other than the reduction in the size

of the neural spine and transverse processes, and the

relative elongation of the centrum, although the length

of the centra remain less than twice their height.

Coracoid (Figure 5). An almost complete left coracoid

is preserved in the Spioenkop assemblage

(BP/1/6113). It is a roughly oval plate with its long

axis measuring 236 mm and oriented subparallel with

the scapula suture and perpendicular to the long axis

of the scapula. The anterior margin is not gently

convex for its entire length but bears a shallow

embayment dorsally. The plate, which is gently convex

laterally and concave medially, is much thinner at the

end furthest from the glenoid (the anterodorsal end).

Figure 7. Right femoral fragments of Eucnemesaurus fortis from Spioenkop. Proximal end and mid section (BP/1/6111) in anterior (A), lateral

(B), posterior (C), and medial (D), views. A, includes cross sections from the distal end of each of the two pieces. Distal end (BP/1/6110) in

anterior (E), lateral (F), posterior (G), and distal (H) views. Scale bar represents 50 mm. Abbreviations: f.c, fibular condyle; f.t, fourth

trochanter; g.t, greater trochanter; i.f, intercondylar fossa; l.c, lateral condyle; l.s, lateral sulcus; m.c, medial condyle, other abbreviations as in

Figure 5.

A. M. Yates8

The coracoid portion of the glenoid is a flattened oval

surface that is slightly turned out so that it can be

viewed laterally but not medially. The coracoid

reaches its maximum mediolateral thickness of

67 mm immediately dorsal to the glenoid. A broad,

indistinct, buttress supports this thickening medially

and the medial opening of the coracoid foramen

pierces the dorsal side of this buttress. This opening

lies close to the scapula suture, whereas on the lateral

side it opens closer to the centre of the bone. The

scapula suture is strongly kinked on the dorsal side of

the medial buttress.

At the posteroventral end of the coracoid, on the

lateral side, there is a prominent laterally projecting

coracoid tubercle. The coracoid tubercle is connected

to the rim of the glenoid by a low, rounded lateroventral

ridge. This ridge and the medial margin of the coracoid

defines a flattened, posteroventrally facing surface.

Scapula (Figure 5). A fragmentary left scapula in

the Spioenkop assemblage articulates perfectly with

the left coracoid, leaving no doubt that they belong to

the same individual. There are two non-contacting

parts: a ventral fragment (BP/1/6114) and a dorsal

fragment (BP/1/6115). The ventral fragment

preserves the glenoid, part of the coracoid articular

surface, the base of the acromion and the base of

Figure 8. Strict consensus tree of 60 most parsimonious trees (tree length ¼ 1094) produced by the analysis of 43 basal dinosaur and

sauropodomorph taxa and 3 outgroup taxa. Taxon labels follow the phylogenetic taxonomy proposed in the text. Arrows represent stem-based

taxa, dots represent node-based taxa.


the scapular blade. The glenoid is a slightly concave

oval surface (78 mm long and 67 mm wide) with its

transverse axis angled slightly with respect to the long

axis of the scapula, so that the glenoid surface is visible

laterally but not medially. A rugose patch of low ridges

and striations on the posteroventral surface, adjacent

to the lip of the glenoid, marks the origin of scapula

head of the m. triceps. The medial side of the base of

the scapula blade shows that there was no ridge or

groove extending parallel to the posteroventral

margin. On the lateral surface there is a broad

shallow fossa, the acromial fossa, above the glenoid

region. The dorsal scapula fragment is too limited to

be of much value. Its width (136 mm) and the

orientation of the bone fibres indicate that like most

sauropodomorphs, the dorsal end was flared.

Pubis (Figures 1a, b, 6a, b). The proximal end of the

left pubis is preserved in TM 119. The proximal body

of the pubis, which bears the iliac articular surface

dorsally, projects anterior to the plane of the pubic

apron. This anterior projection creates a concave

anterior margin, below the iliac articulation, when the

pubis is viewed laterally. The lateral surface of

the proximal body lacks a pubic tubercle. Ventral to

the proximal pubic body is the base of the pubic apron.

It shows that the apron was flattened, transversely

oriented and not much narrower than the distance

between the iliac peduncles in anterior view. Van

Hoepen (1920) also described a distal fragment of the

pubis of TM 119 that could not be located. However

his description makes it clear that there was a

moderate distal swelling that at least as twice as thick

in the anteroposterior dimension than the shaft of the

apron (taken from the minimum measurement on the

proximal pubic fragment).

Femur (Figures 1c, d, 6c, d, 7). The femur is the most

distinctive bone of Eucnemesaurus. Unlike most other

dinosaurs, except Riojasaurus (PVL 3805), a large

rounded tubercle protrudes from the posterior surface

of the proximal end, immediately below the proximal

articular surface. This tubercle is present in TM 119

and NMW 1889-XV-39. It is present as a low mound in

BP/1/6111 but its surface is abraded and the original

tubercle may have been higher. The proximal surface

bears a longitudinal sulcus like most other early

dinosaurs. The medially projecting head is subrounded

in anterior view and lacks a distinct distal edge where

the ligaments of the caput femoris would have been

inserted. It is oriented subhorizontally though it

appears to be distally deflected in NMW 1889-XV-

39. The latter condition is probably the result of the

medial shaft fragment being incorrectly attached to the

head. The proximolateral corner of the femur is

rounded in anterior view. The greater trochanter is a

low, proximodistally oriented, anteriorly projecting

ridge that is located on the lateral margin of the shaft. It

is set some distance from the proximal end of the bone

so that its proximal end is approximately level with the

distal margin of the femoral head. The tall, crest-like

lesser trochanter extends parallel to the greater

trochanter, and the proximodistal axis of the bone,

on the anterior surface of the proximal shaft. It is

situated on the lateral side of the middle of the shaft in

anterior view. It does not extend as far proximal as the

greater trochanter, and its proximal termination is

distal to the distal margin of the femoral head. The

proximal end of the crest is well-preserved in

BP/1/6111 and it shows that the height of the crest

decreases towards the proximal end but the

termination is still abrupt as it is in TM 119. Three

small, flattened spurs with a rounded profile project

proximally from the proximal termination of the lesser

trochanter and suggest that it was the site of a strong

ligamentous attachment. The flat surface between the

lesser trochanter and the greater trochanter bears no

trace of a trochanteric ridge. There is no connection

between the proximal portion of BP/1/6111 and the

segment that bears the fourth trochanter so it is not

possible to say how proximally or distally the fourth

trochanter was placed. However enough of the shaft is

preserved to show that the fourth trochanter cannot be

brought as close to the proximal end as it is in NMW

1889-XV-39. Nevertheless the fourth trochanter itself

is identical in shape to that of the Vienna specimen.

The proximal end of the trochanter is level with

the medial margin of the shaft in posterior view,

the proximal half of the trochanter slopes distolaterally

relative to the long axis of the femoral shaft. After the

midlength of the fourth trochanter its axis curves and

extends approximately parallel to the long axis of the

shaft. The crest also becomes thicker distal to the

midlength. The profile of the fourth trochanter is

rounded rather than the subrectangular shape seen in

most basal sauropodomorphs, except Riojasaurus

(PVL 3805). The distal end of the fourth trochanter

is separated from the femoral shaft by a rounded notch.

Distal to the fourth trochanter the femoral shaft is

slightly wider transversely (92 mm) than it is

anteroposteriorly (83 mm) but this degree of

eccentricity is below that seen in derived anchisaurs

such as Antentonitrus and Melanorosaurus.

The distal end of what is very probably the same

femur (BP/1/6110) was shattered into many pieces

and had washed downstream a short distance. It can

be assembled into a single piece that consists of the

medial, lateral and fibular condyles. A large wedge of

bone is missing from the intercondylar area. The

medial condyle is roughly equivalent in size to the

fibular þ lateral condyles. The distal surfaces of

the medial and fibular condyles are rather flat but

the medial condyle curves proximally at the posterior

end so that this part of the condylar surface faces

A. M. Yates10

posteriorly. There is no extensor depression on the

anterior face of the distal end although the very distal

end, immediately adjacent to the condylar surface is

missing. The surface proximal to the missing wedge is

broad and flattened. A limited, shallow extensor

depression may well have been present at the very

distal end, as it is in NMW 1889-XV-39 (Galton and

Van Heerden 1998), because the anterior margin of

the fibular condyle lies slightly ahead of that of the

medial condyle to the same degree seen in NMW

1889-XV-39. Posteriorly there is a deep fossa between

the medial and lateral condyles. The lateral condyle is

small and sub triangular in distal view, with the apex

pointing posterolaterally. A vague sulcus separates it

from the fibular condyle. This sulcus is continuous

with a deep lateral sulcus that extends proximally.

Tibia (Figures 1e, f, g, h). The tibia of TM 119 could

not be located in the collections of the Transvaal

Museum but it has been well illustrated by Van

Hoepen (1920) and Van Heerden (1979). The

following observations are based on these

illustrations and the measurements given in Van

Hoepen (1920). It is a stout bone where the

maximum length of the proximal articular surface is

42% of the maximum length of the bone. The

cnemial crest is thick and projects forward although

the medial surface curves toward the lateral side. The

fibular condyle is large and indistinct but is clearly

centrally located so that its posterior margin is not

level with the posterior end of the proximal articular

surface. The distal end is transversely wider than it is

anteroposteriorly long. The descending posterolateral

process of the distal tibia extends laterally to draw

level with the anterolateral corner of the distal tibia.

The lateral margin of this process is straight and is

not bevelled or excavated as it is in Plateosauravus

cullingworthi.

Comparison with contemporary taxa

The unusual high diversity of sauropodomorphs in the

lower Elliot Formation warrants detailed comparison

between each taxon and Eucnemesaurus fortis to further

test the validity of this taxon and to aid in the

identification of future specimens.

Plateosauravus cullingworthi differs from Eucne-

mesaurus fortis in the following characteristics: absence

of an accessory lamina in the middle chonos of the

middle dorsal vertebrae; presence of a posteroventral

ridge on the medial side of the scapula; absence of a

large posterior tubercle at the proximal end of the

femur; a low lesser trochanter that is wider than tall; a

more distally placed fourth trochanter (first rise is

1.3–1.4 times the length of the femoral head from

the proximal end); a longitudinally oriented fourth

trochanter (not oblique) that is located away from the

medial edge in caudal view; presence of a bevelled

embayment above the descending posterolateral

process of the distal end of the tibia. If a pubic

fragment that has been tentatively referred to as

Plateosauravus cullingworthi, is correctly identified then

this taxon further differs from Eucnemesaurus fortis in

the presence of a large oval pubic tubercle on the

lateral surface of the proximal end of the pubis.

Melanorosaurus readi differs in the following charac-

teristics: absence of an accessory lamina in the middle

chonos of the middle dorsal vertebrae; narrow hypo-

sphenes that are deeper than the neural canal in the

dorsal vertebrae; proximal caudal vertebrae with

hyposphenal ridges and median ventral fossae;

presence of a posteroventral sulcus on the medial

side of the scapula; a more abbreviated femoral head;

the absence of a large posterior tubercle at the proximal

end of the femur; a lower lesser trochanter that is

placed laterally so that it is visible in posterior view;

a femoral shaft that is close to straight in anterior view;

a moderately eccentric cross-section of the femoral

shaft with the transverse width below the fourth

trochanter exceeding the anteroposterior length; a

subrectangular fourth trochanter without a distal

notch; a longitudinally oriented fourth trochanter

that is situated against the medial edge of the shaft

in posterior view and straddles the midlength of the

femur.

Blikanasaurus cromptoni differs in having a more

robust tibia with a descending posterolateral flange

that lies medial to the lateral margin of the bone in

distal view.

Antetonitrus ingenipes differs in all of the same

femoral characters that differentiate Melanorosaurus

readi, as well as in the following characteristics: tall

dorsal neural arches with slot-shaped neural canals;

deep hyposphenes with median ventral ridges on the

dorsal vertebrae; absence of an accessory lamina in the

middle chonos of the middle dorsal vertebrae; a

descending posterolateral flange of the distal end of

the tibia that lies medial to the lateral margin of the

bone in distal view.

Lastly the unnamed sauropodomorph from Nova

Barletta (Yates 2003) can be distinguished from

Eucnemesaurus fortis by the following characteristics:

low, thick laminae in the dorsal neural arches; absence

of an accessory lamina in the middle chonos of the

middle dorsal vertebrae; absence of a large posterior

tubercle at the proximal end of the femur; a more

abbreviated femoral head; a low lesser trochanter that

is wider than tall; a longitudinally oriented fourth

trochanter (not oblique) that is located away from the

medial edge in caudal view.

Phylogenetic analysis

The sauropodomorph affinity of Eucnemesaurus fortis

and its sister-group relationship with Riojasaurus


incertus is supported by a cladistic analysis of a matrix

of 46 taxa (including 3 outgroup taxa) and 353

characters (see electronic supplemental material for

the character list and character-taxon matrix). The

analysis encompasses a broad range of early dinosaurs

in order to avoid making an a priori decision regarding

the immediate outgroup of Sauropodomorpha. Anal-

ysis of the matrix produces 60 most-parsimonious

trees that are 1094 steps long. The strict consensus of

the 60 fundamental trees is presented here (Figure 7)

but the detailed tree description and robustness tests

of the various nodes will be presented elsewhere.

Eucnemesaurus and Riojasaurus comprise the Rioja-

sauridae in this tree. The Riojasauridae is the basal

branch of a large clade containing all taxa more closely

related to Neosauropoda than to Plateosaurus engel-

hardti. This clade conforms the phylogenetic defi-

nition of Sauropoda that is in current use, i.e. the most

inclusive clade containing Saltasaurus loricatus (a

neosauropod) but not Plateosaurus engelhardti (Wilson

and Sereno 1998). However due to the paraphyly of

the traditional prosauropod assemblage, in this

analysis, and the relatively basal position of Plateo-

saurus engelhardti amongst that assemblage, this

definition ‘captures’ a large number of taxa that

have not been included in the Sauropoda before

(Riojasaurus, Eucnemesaurus, Massospondylus, Color-

adisaurus, Lufengosaurus, Jingshanosaurus and Yunna-

nosaurus) or taxa that are only rarely considered to be

sauropods (Anchisaurus and Melanorosaurus). Thus

the content of Sauropoda has departed radically from

its original concept and it is simplest to re-define

Sauropoda. I propose that Sauropoda be defined as

the most inclusive clade that includes Saltasaurus

loricatus but not Melanorosaurus readi. This definition

is desirable because in phylogenies where the

prosauropod assemblage forms a paraphyletic array,

Sauropoda is still restricted to the clade of specialised

gigantic quadrupeds that form the traditional Sauro-

poda and their closest outgroups whereas in topologies

that have an inclusive, monophyletic Prosauropoda

(e.g. Galton and Upchurch 2004) it makes no

difference whether Plateosaurus or Melanorosaurus

forms the exclusive anchor taxon. One possible

objection to this definition of Sauropoda is that

Melanorosaurus readi is a poorly known taxon and is

likely to be quite labile in phylogenetic hypotheses.

This is no longer the case as there are several high-

quality specimens that can be referred to this taxon

(Galton et al. 2005; Yates 2005). One of these new

specimen includes a complete skull as well as an

articulated manus and pes, arguably making it a

better-known taxon than Saltasaurus loricatus. A new

clade name is now required for the most inclusive

clade containing Saltasaurus loricatus but not Plateo-

saurus engelhardti and I propose that it be called

Massopoda tax. nov.. Note that Massopoda simply

becomes a heterodefinitional synonym of Sauropoda

in phylogenies where Plateosaurus and Melanorosaurus

are included in Prosauropoda.

Discussion

Far from being a basal dinosaur as is usually

suspected, Aliwalia rex, or rather its senior synonym

Eucnemesaurus fortis, is a sauropodomorph and not a

particularly basal one at that. Synapomorphies of

Sauropodomorpha, or less inclusive clades, that are

found in E. fortis include absence of the prezygo-

diapophyseal lamina and its associated anterior

chonos in the middle and posterior dorsal vertebrae,

stout mid-caudal vertebral centra that are less than

twice as long as high (a synapomorphy of sauropodo-

morphs more derived than Saturnalia tupiniquim and

Thecodontosaurus spp.) proximo-distal elongation of

the lesser trochanter (a synapomorphy of sauropodo-

morphs more derived than Saturnalia tupiniquim),

a concave anterior margin of the proximal pubis in

lateral view (a synapomorphy of sauropodomorphs

more derived than Efraasia minor), a distally notched

(semi-pendant) fourth trochanter (a derived character

with an ambiguous distribution that is found in

Riojasauridae and Massospondylidae). In addition

Eucnemesaurus fortis shares some characters uniquely

with Riojasaurus incertus.

Yates (2003) proposed that an unnamed sauropo-

domorph specimen (BP/1/4953) from the lower Elliot

Formation could represent a sister taxon of Rioja-

saurus incertus based on the shared derived presence of

very thick and low laminae of the dorsal neural arches.

However this specimen lacks all of the femoral

synapomorphies of Riojasauridae proposed in this

paper while Eucnemesaurus fortis has ordinary tall, thin

laminae on its dorsal neural arches. Therefore it now

seems rather unlikely that BP/1/4953 shares a close

relationship with Riojasaurus incertus, amongst sauro-

podomorphs. Given the break-up of Prosauropoda

sensu Yates and Kitching (2003) in the analysis

presented here, BP/1/4953 cannot be classified any

further than Sauropodomorpha incertae sedis.

Galton (1985) suggested that an unusual dinosaur-

ian femur, SMNS 51958, from the Norian of Germany

belonged to the same family as ‘Aliwalia rex’. The

specimen consists of a proximal end of a left femur

from the Lowenstein Formation ( ¼ Stubensandstein).

A close relationship between SMNS 51958 and

Eucnemesaurus fortis is unlikely because they differ in

a number of key points. The proximal end of the fourth

trochanter is level with the base of the femoral head

whereas it is far distal to this level in all Eucnemesaurus

fortis specimens including NM 1886-XV-39 where the

fourth trochanter is artificially placed more proximal

than it should be. Furthermore the fourth trochanter

lacks the rounded profile and notched distal end seen in

Eucnemesaurus fortis. The lesser trochanter of SMNS

A. M. Yates12

51958 does not form a proximodistally elongate ridge

as it does in all sauropodomorphs, except Saturnalia

tupiniquim (Langer 2004b). The identity of SMNS

51958 is obscure but it can be confidently excluded

from the clade of Sauropodomorpha more derived than

Saturnalia tupiniquim (including Riojasauridae).

The diversity of sauropodomorphs in the lower

Elliot Formation now increases to six species

(Antetonitrus ingenipes, Blikanasaurus cromptoni, Mela-

norosaurus readi, Plateosauravus cullingworthi, Eucne-

mesaurus fortis and an unnamed sauropodomorph

Yates 2003), making it the richest known Triassic

sauropodomorph fauna. It is frustrating that the

carnivores that utilised this rich resource of prey are so

poorly known. Isolated teeth and undiagnostic

fragments of jaws are not uncommon but no remains

that can conclusively identify them even as far as the

level of Dinosauria, or Rauisuchia, are known. An

isolated tooth crown from the lower Elliot Formation

has been referred to Theropoda (Ray and Chinsamy

2002) but the authors’ reasons for excluding this tooth

from the Rauisuchia were based on comparisons with

just one rauisuchid taxon, Teratosaurus, and another

isolated tooth from the lower Elliot Formation that

cannot be certainly identified. Furthermore the

reported differences between these teeth were the

greater size and subrectangular basal cross section of

the putative theropod tooth. Both of these characters

can vary considerably between species and along a

single jaw, consequently the identification of this tooth

should be considered highly suspect.

Given that Eucnemesaurus is nested deeply within

Sauropodomorpha, it is extremely unlikely that the

maxilla tentatively referred to as Aliwalia rex by Galton

and Van Heerden (1998) belongs to Eucnemesaurus

fortis and the specimen is hereby removed from the

hypodigm of that species. Like other fossils of

carnivorous reptiles from the lower Elliot Formation,

all that can be said of it is that it belongs to a

moderately large archosaur.

Acknowledgements

Emese Bordy, John Hancox and Paul Dirks found

the new Eucnemesaurus specimen while Pepson

Mukanela and Doctor Mbense prepared some of

the bones. I thank Naude Bremmer and Cobus

Visser for allowing us to collect on Spioenkop and

for their hospitality during our stay. Deigo Pol and

Peter Galton are thanked for their review of the

manuscript and their useful comments. I thank

Francis Thackeray for access to specimens in the

Transvaal Museum. Travel to China for the coding

of Chinese taxa used in the phylogenetic analysis was

supported by P.A.S.T.

References

Broom R. 1911. On the dinosaurs of the Stormberg, South Africa.

Ann South African Mus 7:291–308.

Cooper MR. 1980. The first record of the prosauropod dinosaur

Euskelosaurus from Zimbabwe. Arnoldia 9:1–17.

Galton PM, Van Heerden J. 1998. Anatomy of the prosauropod

dinosaur Blikanasaurus cromptoni (Upper Triassic, South Africa),

with notes on other tetrapods from the lower Elliot Formation.

Palaontologische Z 72:163–177.

Galton PM. 1985. The poposaurid thecodontian Teratosaurus

suevicus v. Meyer, plus referred specimens mostly based on

prosauropod dinosaurs, from the Middle Stubensandstein

(Upper Triassic) of Nordwurttemberg. Stuttgart Beitrage zur

Naturkunde (B) 116:1–29.

Galton PM, Upchurch P. 2004. Prosauropoda. In: Weishampel DB,

Dodson P, Osmolska H, editors. The dinosauria. 2nd ed.,

Berkeley: University of California Press. p 232–258.

Galton PM, Van Heerden J, Yates AM. 2005. Postcranial anatomy

of referred specimens of Melanorosaurus from the Upper Triassic

of South Africa. In: Tidwell V, Carpenter K, editors. Thunder-

lizards. The sauropodomorph dinosaurs. Bloomington: Indiana

University Press. p 1–37.

Huene F von. 1906. Ueber die Dinosaurier der aussereuropaischen

Trias. Geol Palaeontologische Abh 8:99–156.

Huxley TH. 1866. On the remains of large dinosaurian reptiles

from the Stormberg Mountains, South Africa. Geol Mag

3:563.

Langer MC. 2004a. Basal saurischians. In: Weishampel DB,

Dodson P, Osmolska H, editors. The dinosauria. 2nd ed.


Langer MC. 2004b. The pelvic and hindlimb anatomy of the stem-

sauropodomorph Saturnalia tupiniquim (Late Triassic, Brazil).

Paleobios 23:1–40.

Novas FE. 1996. Dinosaur monophyly. J Vert Paleontol

16:723–741.

Paul GS. 1988. Predatory dinosaurs of the world. New York: Simon

and Schuster.

Ray S, Chinsamy A. 2002. A theropod tooth from the Late Triassic

of southern Africa. J Biosci 27:295–298.

Seeley HG. 1894. On Euskelosaurus brownii (Huxley). Ann Mag bof

Nat Hist 14:317–340.

Sues H-D. 1990. Staurikosaurus and Herrerasauridae. In: Weisham-

pel DB, Dodson P, Osmolska H, editors. The dinosauria.


Van Heerden J. 1979. The morphology and taxonomy of

Euskelosaurus (Reptilia: Saurischia; Late Triassic) from South

Africa. Navorsinge Van Die Nasionale Mus 4:21–84.

Van Hoepen ECN. 1920. Contributions to the knowledge of the

reptiles of the Karroo Formation. 6. Further dinosaurian

material in the Transvaal Museum. Ann Transvaal Mus

7:7–140.

Wilson JA, Sereno PC. 1998. Early evolution and higher-level

phylogeny of sauropod dinosaurs. Soc Vert Paleontol Memoirs

5:1–68.

Yates AM, Kitching JW. 2003. The earliest known sauropod

dinosaur and the first steps towards sauropod locomotion. Proc

R Soc Lond B 270:1753–1758.

Yates AM. 2003. A definite prosauropod dinosaur from the lower

Elliot Formation (Norian: Upper Triassic) of South Africa.

Palaeontologia africana 39:63–68.

Yates AM. 2005. The skull of the Triassic sauropodomorph

Melanorosaurus readi from South Africa and the definition of

Sauropoda. J Vert Paleontol 25 (supplement to 3):132A.


Character List

1. Skull to femur ratio: greater than (0), or less than

(1), 0.6 (modified from Gauthier 1986).

2. Lateral plates appressed to the labial side of the

premaxillary, maxillary and dentary teeth: absent

(0) or present (1) (Upchurch 1995).

3. Relative height of the rostrum at the posterior

margin of the naris: more than (0), or less than

(1), 0.6 of the height of the skull at the middle of

the orbit (Langer 2004).

4. Foramen in the middle of the lateral surface

of the premaxillary body: absent (0), or present

(1).

5. Distal end of the dorsal premaxillary process:

tapered (0) or transversely expanded (1) (Sereno

1999).

6. Profile of premaxilla: convex (0) or with an

inflection at the base of the dorsal process (1)

(Upchurch 1995).

7. Size and position of the posterolateral process

of premaxilla: large and lateral to the anterior

process of the maxilla (0) or small and medial to

the anterior process of the maxilla (1).

8. Relationship between posterolateral process of

the premaxilla and the anteroventral process of

the nasal: broad sutured contact (0), point

contact (1) or separated by maxilla (2) (modified

from Gauthier 1986). Ordered.

9. Posteromedial process of the premaxilla: absent

(0) or present (1) (Rauhut 2003).

10. Shape of the anteromedial process of the maxilla:

narrow, elongated and projecting anterior to

lateral premaxilla-maxilla suture (0), short, broad

and level with lateral premaxilla-maxilla suture

(1).

11. Development of external narial fossa: absent to

weak (0) or well developed with sharp posterior

and anteroventral rims (1).

12. Size and position of subnarial foramen: absent

(0), small (no larger than adjacent maxillary

neurovascular foramina) and positioned outside

of narial fossa (1), or large and on the rim of, or

inside, the narial fossa (2) (modified from Sereno

et al. 1993). Ordered.

13. Shape of subnarial foramen: rounded (0) or slot-

shaped (1).

14. Maxillary contribution to the margin of the narial

fossa: absent (0) or present (1).

15. Diameter of external naris: less than (0), or

greater than (1), 0.5 of the orbital diameter

(Wilson and Sereno 1998).

16. Shape of the external naris (in adults): rounded

(0) or subtriangular with an acute posteroventral

corner (1) (Galton and Upchurch 2004).

17. Level of the anterior margin of the external naris:

anterior to (0), or posterior to (1), the midlength

of the premaxillary body (Rauhut 2003).

18. Level of the posterior margin of external naris:

anterior to, or level with the premaxilla-maxilla

suture (0), posterior to the first maxillary

alveolus (1) or posterior to the midlength of the

maxillary tooth row and the anterior margin of

the antorbital fenestra (2) (modified from Wilson

and Sereno 1998). Ordered.

19. Dorsal profile of the snout: straight to gently

convex (0) or with a depression behind the

naris (1).

20. Elongate median nasal depression: absent (0) or

present (1) (Sereno 1999).

21. Width of anteroventral process of nasal at its

base: less than (0) or greater than (1) width of

anterodorsal process at its base (modified from

Sereno 1999).

22. Nasal relationship with dorsal margin of ant-

orbital fossa: not contributing to the margin of

the anorbital fossa (0), lateral margin overhangs

the antorbital fossa and forms its dorsal margin

(1), overhang extensive, obscuring the dorsal

lacrimal-maxilla contact in lateral view (2)

(modified from Sereno 1999).

23. Pointed caudolateral process of the nasal

overlapping the lacrimal: absent (0) or present

(1) (Sereno 1999).

24. Anterior profile of the maxilla: slopes continu-

ously towards the rostral tip (0) or with a strong

inflection at the base of the ascending ramus,

creating a rostral ramus with parallel dorsal and

ventral margins (1) (Sereno et al. 1996).

25. Length of rostral ramus of the maxilla: less than

(0), or greater than (1), its dorsoventral depth

(Sereno et al. 1996).

26. Shape of the main body of the maxilla: tapering

posteriorly (0) or dorsal and ventral margins

parallel for most of their length (1).

27. Shape of the ascending ramus of the maxilla

in lateral view: tapering dorsally (0) or with an

anteroposterior expansion at the dorsal end (1)

28. Rostrocaudal length of the antorbital fossa:

greater (0), or less (1), than that of the orbit

(Yates 2003a).

29. Posteroventral extent of medial wall of antorbital

fossa: reaching (0), or terminating anterior to

(1), the anterior tip of the jugal (modified from

Galton and Upchuch 2004).

30. Development of the antorbital fossa on the

ascending ramus of the maxilla: deeply

impressed and delimited by a sharp, scarp-like

rim (0), weakly impressed and delimited by a

rounded rim or a change in slope (1).

31. Shape of the rostral margin of the antorbital

fenestra: strongly concave, roughly parallel to the

rostral margin of the antorbital fossa, creating a

narrow antorbital fossa (0) or straight to gently

concave creating a broad, subtriangular ant-

orbital fossa (1) (Galton 1985a).

A. M. Yates14

32. Size of the neurovascular foramen at the caudal

end of the lateral maxillary row: not larger than

the others (0) or distinctly larger than the others

in the row (1) (Yates 2003a).

33. Direction that the neurovascular foramen at the

caudal end of the lateral maxillary row opens:

caudally (0), rostrally, ventrally or laterally (1)


34. Arrangement of lateral maxillary neurovascular

foramina: linear (0) or irregular (1) (modified

from Sereno 1999).

35. Dorsal exposure of the lacrimal: present (0) or

absent (1) (Gauthier 1986).

36. Shape of the lacrimal: dorsoventrally short and

block-shaped (0) or dorsoventrally elongate and

shaped like an inverted ‘L’(1) (Rauhut 2003).

37. Orientation of the lacrimal orbital margin:

strongly sloping anterodorsally (0) or erect and

close to vertical (1).

38. Length of the anterior ramus of the lacrimal:

greater than (0), or less than (1), half the length

of the ventral ramus, or absent altogether (2)

(modified from Galton 1990). Ordered.

39. Web of bone spanning junction between anterior

and ventral rami of lacrimal: absent and

antorbital fossa laterally exposed (0) or present,

obscuring posterodorsal corner of antorbital

fossa (1).

40. Extension of the antorbital fossa onto the ventral

end of the lacrimal: present (0) or absent (1)

(modified from Wilson and Sereno 1998).

41. Length of the caudal process of the prefrontal:

short (0), or elongated (1), so that total

prefrontal length is equal to the rostrocaudal

diameter of the orbit (Galton 1985a).

42. Ventral process of prefrontal extending down the

posteromedial side of the lacrimal: present (0) or

absent (1) (Wilson and Sereno 1998).

43. Maximum transverse width of the prefrontal: less

than (0), or more than (1), 0.25 of the skull

width at that level (modified from Galton 1990).

44. Shape of the orbit: subcircular (0) or ventrally

constricted making the orbit subtriangular (1)


45. Slender anterior process of the frontal intruding

between the prefrontal and the nasal: absent (0)

or present (1) (modified from Sereno 1999).

46. Jugal-lacrimal relationship: lacrimal overlapping

lateral surface of jugal or abutting it dorsally (0),

or jugal overlapping lacrimal laterally (1) (Sereno

et al. 1993).

47. Shape of the suborbital region of the jugal: an

anteroposteriorly elongate bar (0) or an antero-

posteriorly shortened plate (1).

48. Jugal contribution to the antorbital fenestra:

absent (0) or present (1) (Holtz 1994).

49. Dorsal process of the anterior jugal: present (0)

or absent (1) (modified from Rauhut 2003).

50. Ratio of the minimum depth of the jugal below

the orbit to the distance between the rostral

end of the jugal and the rostroventral corner of

the lower temporal fenestra: less than (0), or

greater than (1), 0.2 (modified from Galton

1985a).

51. Transverse width of the ventral ramus of the

postorbital: less than (0), or greater than (1), its

rostrocaudal width at mid shaft (Wilson and

Sereno 1998).

52. Shape of the dorsal margin of postorbital in

lateral view: straight to gently curved (0) or with

a distinct embayment between the anterior and

posterior dorsal processes (1).

53. Height of the postorbital rim of the orbit: flush

with the posterior lateral process of the

postorbital (0) or raised so that it projects

laterally to the posterior dorsal process (1).

54. Postfrontal bone: present (0) absent (1) (Sereno

et al. 1993).

55. Position of the rostral margin of the lower

temporal fenestra: behind the orbit (0), extends

under the rear half of the orbit (1) or extends as

far forward as the midlength of the orbit (2)

(modified from Upchurch 1995). Ordered.

56. Frontal contribution to the supratemporal

fenestra: present (0) or absent (1) (modified

from Gauthier 1986).

57. Orientation of the long axis of the supratemporal

fenestra: longitudinal (0) or transverse (1)


58. Medial margin of supratemporal fossa: simple

smooth curve (0) or with a projection at the

frontal/postorbital-parietal suture producing a

scalloped margin (1) (Leal et al. 2004).

59. Length of the quadratojugal ramus of the

squamosal relative to the width at its base: less

than (0) or greater than (1) four times its width

(Sereno 1999).

60. Proportion of lower temporal fenestra bordered

by squamosal: more than (0), or less than (1), 0.5

of the depth of the lower temporal fenestra.

61. Squamosal-quadratojugal contact: present (0) or


62. Angle of divergence between jugal and squa-

mosal rami of quadratojugal: close to 908 (0) or

close to parallel (1).

63. Length of jugal ramus of quadratojugal: no

longer than (0), or longer than (1), the

squamosal ramus (Wilson and Sereno 1998).

64. Shape of the rostral end of the jugal ramus of

the quadratojugal: tapered (0) or dorsoventrally

expanded (1) (Wilson and Sereno 1998).

65. Relationship of quadratojugal to jugal: jugal

overlaps the lateral surface of the quadratojugal

(0), quadratojugal overlaps the lateral surface of

the jugal (1), or quadratojugal sutures along the

ventrolateral margin of the jugal (2).


66. Position of the quadrate foramen: on the

quadrate-quadratojugal suture (0) or deeply

incised into, and partly encircled by, the

quadrate (1) (Rauhut 2003).

67. Shape of posterolateral margin of quadrate:

sloping anterolaterally from posteromedial ridge

(0), everted posteriorly creating a posteriorly

facing fossa (1), posterior fossa deeply excavated,

invading quadrate body (2) (Wilson and Sereno

1998). Ordered.

68. Exposure of the lateral surface of the quadrate

head: absent, covered by lateral sheet of the

squamosal (0) or present (1) (Sereno et al.

1993).

69. Proportion of the length of the quadrate that is

occupied by the pterygoid wing: at least 70% (0)

or greater than 70% (1) (Yates 2003a).

70. Depth of the occipital wing of the parietal: less

than (0), or more than 1.5 (1) times the depth of

the foramen magnum (Wilson and Sereno

1998).

71. Position of vena capitis foramina on occiput:

between supraoccipital and parietal (0) or on the

supraoccipital (1).

72. Dorsal fenestra between supraoccipital and

parietals: absent (0) or present (1).

73. Shape of the supraoccipital: diamond-shaped,

at least as high as wide (0), or semilunate and

wider than high (1) (Yates 2003b).

74. Orientation of the supraoccipital plate: erect to

gently sloping (0) or strongly sloping forward so

that the dorsal tip lies level with the basipterygoid

processes (1) (Galton and Upchurch 2004).

75. Orientation of the paroccipital processes: slightly

dorsolaterally directed to horizontal (0), or

ventrolaterally directed (1) in occipital view

(Rauhut 2003).

76. Size of the posttemporal fenestra: large fenestra

(0) small hole that is no larger than any of the

foramina for the cranial nerves (1).

77. Exit of the mid-cerebral vein: through trigeminal

foramen (0) or through a separate foramen

anterodorsal to trigeminal foramen (1) (Rauhut

2003).

78. Shape of the floor of the braincase in lateral

view: relatively straight with the basal tuberae,

basipterygoid processes and parasphenoid ros-

trum roughly aligned (0), bent with the

basipterygoid processes and the parasphenoid

rostrum below the level of the basioccipital

condyle and the basal tuberae (1) or bent with

the basal tuberae lowered below the level of the

basioccipital and the parasphenoid rostrum

raised above it (2) (modified from Galton

1990). Unordered.

79. Shape of basal tuberae: knob-like, with basis-

penoidal component rostral to basioccipital

component (0), or forming a transverse ridge

with the basisphenoidal component lateral to the

basioccipital component (1).

80. Length of the basipterygoid processes (from the

top of the parasphenoid to the tip of the process):

less than (0), or greater than (1), the height of the

braincase (from the top of the parasphenoid to the

top of the supraoccipital) (Benton et al. 2000).

81. Ridge formed along the junction of the parabasi-

sphenoid and the basioccipital, between the basal

tuberae: present with a smooth rostral face (0),

present with a median fossa on the rostral face (1),

or absent with the basal tuberae being separated by

a deep caudally opening U-shaped fossa (2).

Unordered.

82. Deep septum spanning the interbasipterygoid

space: absent (0) or present (1) (Galton 1990).

83. Dorsoventral depth of the parashenoid rostrum:

much less than (0) or about equal to the transverse

width (1) (Yates 2003a).

84. Shape of jugal process of ectopterygoid: gently

curved (0) or strongly recurved and hook-like (1)

(Yates 2003a).

85. Pneumatic fossa on the ventral surface of the

ectopterygoid: present (0) or absent (1) (Sereno

et al. 1996).

86. Relationship of the ectopterygoid to the pterygoid:

ectopterygoid overlapping the ventral (0), or

dorsal (1), surface of the pterygoid (Sereno et al.

1993).

87. Position of the maxillary articular surface of the

palatine: along the lateral margin of the bone (0)

or at the end of a narrow anterolateral process (1)


88. Centrally located tubercle on the ventral surface

of palatine: absent (0) or present (1).

89. Medial process of the pterygoid forming a hook

around the basipterygoid process: absent (0), flat

and blunt-ended (1) or bent upwards and

pointed (2) (modified from Wilson and Sereno

1998). Ordered.

90. Length of the vomers: less than (0), or more than

(1), 0.25 of the total skull length.

91. Position of jaw joint: no lower than the level of

the dorsal margin of the dentary (0) or depressed

well below this level (1) (Sereno 1999).

92. Shape of upper jaws in ventral view: narrow with

an acute rostral apex (0) or broad and U-shaped

(1) (Wilson and Sereno 1998).

93. Length of the external mandibular fenestra:

more than (0), or less than (1), 0.1 of the length

of the mandible.

94. Caudal end of dentary tooth row medially inset

with a thick lateral ridge on the dentary forming

a buccal emargination: absent (0) or present (1)

(Gauthier 1986).

95. Height: length ratio of the dentary: less than (0),

or greater than (1), 0.2 (modified from Benton

et al. 2000).

A. M. Yates16

96. Orientation of the symphyseal end of the dentary:

in line with the long axis of the dentary (0) or

strongly curved ventrally (1) (Sereno 1999).

97. Position of first dentary tooth: adjacent to

symphysis (0) or inset one tooth’s width from

the symphysis (1) (Sereno 1999).

98. Dorsoventral expansion at the symphyseal end of

the dentary: absent (0) or present (1) (Wilson

and Sereno 1998).

99. Splenial foramen: absent (0), present and

enclosed (1), or present and open anteriorly (2)

(Rauhut 2003). Ordered.

100. Splenial-angular joint: flattened sutured contact

(0), synovial joint surface between tongue-like

process of angular fitting in groove of the splenial

(1) (Sereno et al. 1993).

101. A stout, triangular, medial process of the

articular, behind the glenoid: present (0) or

absent (1) (Yates 2003a).

102. Length of the retroarticular process: less than

(0), or greater than (1), than the depth of the

mandible below the glenoid (Yates 2003a).

103. Strong medial embayment behind glenoid of the

articular in dorsal view: absent (0), or present (1)

(Yates and Kitching 2003).

104. Number of premaxillary teeth: four (0) or more

than four (1) (Galton 1990).

105. Number of dentary teeth (in adults): less than 18

(0), 18 or more (1) (modified from Wilson and

Sereno 1998).

106. Arrangement of teeth within the jaws: linearly

placed, crowns not overlapping (0), or imbri-

cated with distal side of tooth overlapping mesial

side of the succeeding tooth (1).

107. Orientation of the maxillary tooth crowns: erect

(0) or procumbent (1) (modified Gauthier

1986).

108. Orientation of the dentary tooth crowns: erect

(0) or procumbent (1) (modified Gauthier

1986).

109. Teeth with basally constricted crowns: absent (0)

or present (1) (Gauthier 1986).

110. Tooth – tooth occlusal wear facets: absent (0) or

present (1) (Wilson and Sereno 1998).

111. Mesial and distal serrations of the teeth: fine and

set at right angles to the margin of the tooth (0)

or coarse and angled upwards at an angle of 458

to the margin of the tooth (1) (Benton et al.

2000).

112. Distribution of serrations on the maxillary and

dentary teeth: present on both the mesial and

distal carinae (0), absent on the posterior carinae

(1), absent on both carinae (2) (Wilson 2002).

Unordered.

113. Long axis of the tooth crowns distally recurved:

present (0) or absent (1) (Gauthier 1986).

114. Texture of the enamel surface: entirely smooth

(0), finely wrinkled in some patches (1), or

extensively and coarsely wrinkled (2) (modified

from Wilson and Sereno 1998).

115. Lingual concavities of the teeth: absent (0) or

present (1) (Upchurch 1995).

116. Longitudinal labial grooves on the teeth: absent


117. Distribution of the serrations along the mesial

and distal carinae of the tooth: extend along

most of the length of the crown (0) or are

restricted to the upper half of the crown (1)

(Yates 2003a).

118. Number of cervical vertebrae: 8 or fewer (0), 9 to

10 (1), 12–13 (2) or more than 13 (3) (modified

from Wilson and Sereno 1998). Ordered.

119. Shallow, dorsally facing fossa on the atlantal

neurapophysis bordered by a dorsally everted

lateral margin: absent (0) or present (1) (Yates

and Kitching 2003).

120. Width of axial intercentrum: less than (0), or

greater than (1), width of axial centrum (Sereno

1999).

121. Position of axial prezgapophyses: on the antero-

lateral surface of the neural arch (0) or mounted

on anteriorly projecting pedicels (1).

122. Posterior margin of the axial postzygapophyses:

overhang the axial centrum (0) or are flush with

the caudal face of the axial centrum (1) (Sereno

1999).

123. Length of the axial centrum: less than (0), or at

least (1), 3 times the height of the centrum.

124. Length of the anterior cervical centra (ce 3–5):

no more than (0), or greater than (1), the length

of the axial centrum.

125. Length of middle to posterior cervical centra

(ce 6–8): no more than (0), or greater than (1),

the length of the axial centrum.

126. Dorsal excavation of the cervical parapophyses:

absent (0) or present (1) (Upchurch 1998).

127. Lateral compression of the anterior cervical

vertebrae: centra are no higher than they are

wide (0) or are approximately 1.25 times higher

than wide (1) (Upchurch 1998).

128. Relative elongation of the anterior cervical centra

(ce 3–5): lengths of the centra are less than 2.5

times the height of their anterior faces (0),

lengths are between 2.5 and 4 times the height

of their anterior faces (1) or the length of at least

ce 4 or 5 exceed 4 times the anterior centrum

height (2) (modified from Sereno 1999).

Ordered.

129. Ventral keels on cranial cervical centra: present (0)

or absent (1) (modified from Upchurch 1998).

130. Height of the mid cervical neural arches: no

more than (0), or greater than (1), height of the

posterior centrum face.

131. Cervical epipophyses on the dorsal surface of the

postzygapophyses: absent (0), or present (1), on

at least some cervical vertebrae.


132. Caudal ends of cranial, postaxial epipophyses:

with a free pointed tip (0) or joined to the

postzygapophysis along their entire length (1).

133. Shape of the epipophyses: tall ridges (0) or

flattened, horizontal plates (1) (Yates 2003a).

134. Epipophyses overhanging the rear margin of the

postzygapophyses: absent (0), or present (1), in

at least some postaxial cervical vertebrae (Sereno

et al. 1993).

135. Anterior spur-like projections on mid-cervical

neural spines: absent (0) or present (1).

136. Shape of mid-cervical neural spines: less than

(0), or at least (1), twice as long as high.

137. Shape of cervical rib shafts: short and postero-

ventrally directed (0) or longer than the length of

their centra and extending parallel to cervical

column (1) (Sereno 1999).

138. Position of the base of the cervical rib shaft: level

with, or higher than the ventral margin of the

cervical centrum (0) or located below the ventral

margin due to a ventrally extended parapophysis


139. Postzygadiapophyseal lamina in cervical neural

arches 4–8: present (0) or absent (1) (Yates

2003a).

140. Laminae of the cervical neural arches 4 to 8:

well developed tall laminae (0) or weakly

developed low ridges (1) (Wilson and Sereno

1998).

141. Shape of anterior centrum face in cervical

centra: concave (0), flat (1) or convex (2)

(modified from Gauthier 1986). Ordered.

142. Ventral surface of the centra in the cervico-dorsal

transition: transversely rounded (0) or with

longitudinal keels (1) (Rauhut 2003).

143. Number of vertebrae between cervicodorsal

transition and primordial sacral vertebrae: 15

to 16 (0) or no more than 14 (1) (modified from

Wilson and Sereno 1998).

144. Lateral surfaces of the dorsal centra: with at

most a vague, shallow depressions (0), with

deep fossae that approach the midline (1) or

with invasive, sharp-rimmed pleurocoels (2)

(Gauthier 1986). Ordered.

145. Oblique ridge dividing pleural fossa of cervical

vertebrae: absent (0) or present (1) (Wilson and

Sereno 1998).

146. Laterally expanded tables at the midlength of the

dorsal surface of the neural spines: absent in all

vertebrae (0), present on the pectoral vertebrae

(1) or present on the pectoral and cervical

vertebrae (2) (Yates and Kitching 2003).

Ordered.

147. Dorsal centra: entirely amphicoelous to amphi-

platyan (0) first two dorsals are opisthocoelous

(1), or cranial half of dorsal column is

opisthocoelous (2) (Wilson and Sereno 1998).

Ordered.

148. Shape of the posterior dorsal centra: relatively

elongated for their size (0) strongly axially

compressed for their size (1) (modified from

Novas 1993).

Various authors have noted that the posterior

dorsal vertebrae of some dinosaur taxa, notably

Herrerasaurus, are much shorter, than they are

tall. This is contrast to the condition in basal

dinosauromorphs (e.g. Marasuchus lilloensis) and

most basal dinosaurs (e.g. Lesothosaurus diagno-

sticus; Dilophosaurus wetherilli; Thecodontosaurus

antiquus). Thus the shortening of the posterior

dorsal vertebral centra, so that the length:height

ratio is less than 1.0 has been used as a character

in several cladistic analyses. Conversely some

analyses take a roughly equant centrum to be the

primitive condition and describe the elongation

of the posterior dorsal centra as derived.

Regardless of the polarity of this character,

these analyses fail to take into account the

allometric changes that occur in the posterior

dorsal centra with increasing body size. If the

length of the posterior dorsal centra of various

sized basal sauropodomorphs are plotted against

their elongation index (EI¼ centrum length/

posterior centrum height) we can see that they

plot close to a line with a negative slope. The very

largest basal sauropodomorphs have short

posterior dorsal vertebrae with EI’s of less than

1 where as small ones approach 1.5. So it would

seem that EI is dependant upon size and that

simple variation in EI should not be used to

discriminate different evolutionary states. How-

ever it is clear that there is variation amongst

the EI of dinosauromorphs that cannot be

accounted for by size alone. When a range of

dinosauromorphs are added to the plot, most

form a linear cloud that is centred upon the

regression line for the basal sauropodomorph

data. Early neotheropods like Lilliensternus and

Dilophosaurus tend to plot above the line,

indicating that they have slightly elongated

vertebrae, for their size, compared to other

early dinosauromorphs. However this variation

is not pronounced, as they still fall within the

bounds of the main cloud and either reflect a

derived condition within Neotheropoda or a

synapomorphy of the group. In either case this

variation is not parsimony informative for the

present analysis and it is ignored. On the other

hand, Herrerasaurus and Staurikosaurus plot well

below the main dinosauromorph cloud and are

separated by a distinct gap in morphospace.

There it is clear that, for their size they have

posterior dorsal centra that are markedly

compressed in the anteroposterior dimension

and they are coded as having a different

character state from other dinosauromorphs.

A. M. Yates18

149. Laminae bounding triangular infradiapophyseal

fossae (chonae) on dorsal neural arches: absent

(0) or present (1) (Wilson 1999).

150. Location of parapophysis in first two dorsals: at

the anterior end of the centrum (0), or located at

the midlength of the centrum, within the middle

chonos (1).

151. Parapophyses of the dorsal column completely

shift from the centrum to the neural arch:

anterior (0), or posterior (1) to the 13th presacral

vertebra (Langer 2004).

152. Orientation of the transverse processes of the

dorsal vertebrae: most horizontally directed (0)

or all upwardly directed (1) (Upchurch 1998).

153. Contribution of the paradiapophyseal lamina to

the margin of the anterior chonos in mid dorsal

vertebrae: present (0) or prevented by high

placement of parapophysis.

154. Hyposphenes in the dorsal vertebrae: absent (0),

present but less than the height of the neural

canal (1), or present and equal to the height of

the neural canal (2) (modified from Gauthier

1986). Ordered.

155. Prezygadiapophyseal lamina and associated

anterior triangular fossa (chonos): present on

all dorsals (0) or absent in mid dorsals (1) (Yates

2003a).

156. Anterior centroparapophyseal lamina in dorsal

vertebrae: absent (0) or present (1) (Wilson

2002).

157. Prezygaparapophyseal lamina in dorsal verte-

brae: absent (0) or present (1).

158. Accessory lamina dividing posterior chonos from

postzygapophysis: absent (0) or present (1).

159. Lateral pneumatic fenestra in middle chonos of

middle and posterior dorsal vertebrae opening

into neural cavity: absent (0) or present (1)


160. Separation of lateral surfaces of anterior dorsal

neural arches under transverse processes: widely

spaced (0) or only separated by a thin midline

septum (1) (Upchurch et al. 2004).

161. Height of dorsal neural arches, from neurocen-

tral suture to level of zygapophyseal facets: much

less than (0), subequal to, or greater than (1),

height of centrum.

162. Height of the dorsal neural spines: greater than

(0), or less than (1), 1.5 times the length of the

base of the spine (modified from Bonaparte

1986).

163. Shape of posterior dorsal neural canal: sub-

circular (0) or slit-shaped (1) (Wilson and

Sereno 1998).

164. Height of middle dorsal neural spines: less than

the length of the base (0), higher than the length

of the base but less than 1.5 times the length of

the base (1) or greater than 1.5 times the length

of the base (2).

165. Shape of anterior dorsal neural spines: lateral

margins parallel in anterior view (0) or transver-

sely expanding towards dorsal end (1).

166. Cross sectional shaped of dorsal neural spines:

transversely compressed (0), broad and triangu-

lar (1) or square-shaped in posterior vertebrae

(2) (modified from Bonaparte 1986).

167. Spinodiapophyseal lamina on dorsal vertebrae:

absent (0), present and separated from spino-

postzygapophyseal lamina (1) or present and

joining spinopostzygapophyseal lamina to create

a composite posterolateral spinal lamina (Wilson

and Sereno 1998).

168. Well developed, sheet-like suprapostzygapophy-

seal laminae: absent (0), present on at least the

caudal dorsal vertebrae (1) (Bonaparte 1986).

169. Shape of the spinopostzygapophyseal lamina in

middle and posterior dorsal vertebrae: singular

(0) or bifurcated at its distal end (1) (Wilson

2002).

170. Shape of posterior margin of middle dorsal

neural spines in lateral view: approximately

straight (0) or concave with a projecting postero-

dorsal corner (1) (Yates 2003c).

171. Transversely expanded plate-like summits of

posterior dorsal neural spines: absent (0), or

present (1) (Novas 1993).

172. Last presacral rib: free (0) or fused to vertebra (1).

173. Sacral rib much narrower than the transverse

process of the first primordial sacral vertebra

(and dorso-sacral if present) in dorsal view:

absent (0) or present (1) (Yates and Kitching

2003).

174. Number of dorsosacral vertebrae: none (0), one

(1) or two (2) (modified Gauthier 1986).

175. Caudosacral vertebra: absent (0) or present (1)

(Galton and Upchurch 2004).

176. Shape of the iliac articular facets of the first

primordial sacral rib: singular (0) or divided into

dorsal and ventral facets separated by a non-

articulating gap (1).

177. Depth of the iliac articular surface of the

primordial sacrals: less than (0), or greater than

(1), 0.75 of the depth of the ilium (modified from

Novas, 1992).

178. Sacral ribs contributing to the rim of the

acetabulum: absent (0) or present (1) (Wilson

2002).

179. Posterior and anterior expansion of the trans-

verse processes of the first and second primordial

sacral vertebrae, respectively, partly roofing the

intercostals space: absent (0) or present (1)

(Langer 2004).

180. Length of first caudal centrum: greater than (0),

or less than (1), its height (Yates 2003a).

181. Length of base of the proximal caudal neural

spines: less than (0), or greater than (1), half the

length of the neural arch (Gauthier 1986).


182. Position of postzygapophyses in proximal caudal

vertebrae: protruding with an interpostzygapo-

physeal notch visible in dorsal view (0) or placed

on either side of the caudal end of the base of the

neural spine without any interpostzygapophyseal

notch (1) (Yates 2003a).

183. A hyposphenal ridge on caudal vertebrae: absent


184. Depth of the bases of the proximal caudal

transverse processes: shallow, restricted to the

neural arches (0), deep extending from the

centrum to the neural arch (1) (Upchurch

1998).

185. Position of last caudal vertebra with a protruding

transverse process: distal (0) or proximal (1) to

caudal 16 (Wilson 2002).

186. Orientation of posterior margin of proximal

caudal neural spines: sloping posterodorsally (0)

or vertical (1) (Novas, 1992).

187. Longitudinal ventral sulcus on proximal and

middle caudal vertebrae: present (0) or absent

(1) (modified from Upchurch 1995).

188. Length of midcaudal centra: greater than (0), or

less than (1), twice the height of their anterior

faces (Yates 2003a).

189. Cross-sectional shape of the distal caudal centra:

oval with rounded lateral and ventral sides (0) or

square-shaped with flattened lateral and ventral

sides (1).

190. Length of distal caudal prezygapophyses: short,

not overlapping the preceding centrum by more

than a quarter (0) or long and overlapping the

preceding the centrum by more than a quarter

(Gauthier 1986).

191. Shape of the terminal caudal vertebrae: unfused,

size decreasing toward tip (0) or expanded and

fused to form a club-shaped tail (1) (Upchurch

1995).

192. Length of the longest chevron: is less than (0),

or greater than (1), twice the length of the

preceding centrum (modified from Yates 2003a).

193. Anteroventral process on distal chevrons: absent


194. Midcaudal chevrons with a ventral slit: absent


195. Longitudinal ridge on the dorsal surface of the

sternal plate: absent (0) or present (1)

(Upchurch 1998).

196. Craniocaudal length of the acromion process of

the scapula: less than (0), or greater than (1), one

and a half times the minimum width of the

scapula blade (Wilson and Sereno 1998).

197. Minimum width of the scapula: is less than (0),

or greater than (1), 20% of its length (Gauthier

1986).

198. Caudal margin of the acromion process of the

scapula: rises from the blade at angle that is less

than (0), or greater than (1), 658 from the long

axis of the scapula, at its steepest point (modified

from Novas, 1992).

199. Flat caudoventrally facing surface on the

coracoid between glenoid and coracoid tubercle:

absent (0) or present (1) (Yates and Kitching

2003).

200. Coracoid tubercle: present (0) or absent (1)

(modified from Perez-Moreno et al. 1994).

201. Length of the humerus: less than 55% (0), 55–

65% (1), 65–70% (2), or more than 70% (3), of

the length of the femur (modified from Gauthier

1986). Ordered.

202. Shape of the deltopectoral crest: subtriangular

(0) or subrectangular (1) (Gauthier 1986).

203. Length of the deltopectoral crest of the humerus:

less than 30% (0), 30–50% (1), or greater than

50% (2), of the length of the humerus (modified

from Sereno et al. 1993). Ordered.

204. Shape of the anterolateral margin of the

deltopectoral crest of the humerus: straight (0)

or strongly sinuous (1) (Yates 2003a).

205. Rugose pit centrally located on the lateral surface

of the deltopectoral crest: absent (0) or present

(1).

206. Well-defined fossa on the distal flexor surface of

the humerus: present (0) or absent (1) (Yates

and Kitching 2003).

207. Transverse width of the distal humerus: is less

than (0), or greater than (1), 33% of the length of

the humerus (Langer 2004).

208. Shape of the entepicondyle of the distal

humerus: rounded process (0) or with a flat

distomedially facing surface bounded by a sharp

proximal margin (1).

209. Length of the radius: greater than (0), or less

than (1), 80% of the humerus (Langer 2004).

210. Deep radial fossa, bounded by an anterolateral

process, on proximal ulna: absent (0) or present


211. Olecranon process on proximal ulna: present (0)

or absent (1) (Wilson and Sereno 1998).

212. Maximum linear dimensions of the ulnare and

radiale: exceed that of at least one of the first

three distal carpals (0) or are less than any the

distal carpals (1) (Yates 2003a).

213. Transverse width of the first distal carpal: less

than (0), or greater than (1), 120% of the

transverse width of the second distal carpal

(Sereno 1999).

214. Sulcus across the medial end of the first distal

carpal: absent (0) or present (1).

215. Lateral end of first distal carpal: abuts (0), or

overlaps (1), second distal carpal (Yates 2003a).

216. Second distal carpal: does (0), or does not (1),

completely cover the proximal end of the second

metacarpal (Yates and Kitching 2003).

217. Ossification of the fifth distal carpal: present (0)

or absent (1).

A. M. Yates20

218. Length of the manus: less than 38% (0), between

38 and 45% (1), or greater than 45% (2), of the

humerus þ radius (modified from Sereno et al.

1993). Ordered.

219. Shape of metacarpus: flattened to gently curved

and spreading (0) or a colonnade of subparallel

metacarpals tightly curved into a ‘u’ shape (1)


220. Proximal width of first metacarpal: less than (0),

or greater than (1), the proximal width of the

second metacarpal (modified from Gauthier

1986).

221. Minimum transverse shaft width of first meta-

carpal: less than (0), or greater than (1), twice

the minimum transverse shaft width of second

metacarpal.

222. Proximal end of first metacarpal: flush with other

metacarpals (0) or inset into the carpus (1)

(Sereno 1999).

223. Shape of the first metacarpal: proximal width less

than 65% (0), between 65% and 80% (1),

between 80% and 100% (2), or greater than

100% (3), of its length (modified from Sereno

1999). Ordered.

224. Strong assymetry in the lateral and medial distal

condyles of the first metacarpal: absent (0) or

present (1) (Gauthier 1986).

225. Deep distal extensor pits on the second and third

metacarpals: absent (0) or present (1) (Novas

1993).

226. Shape of the distal ends of second and third

metacarpals: subrectangular in distal view (0)

or trapezoidal with flexor rims of distal

collateral ligament pits flaring beyond extensor

rims (1).

227. Shape of the fifth metacarpal: longer than wide at

the proximal end with a flat proximal surface (0)

or close to as wide as it is long with a strongly

convex proximal articulation surface (1) (Yates

2003a).

228. Length of the fifth metacarpal: less than (0), or

greater than (1), 75% of the length of the third

metacarpal (Upchurch 1998).

229. Length of manual digit one: less than (0), or

greater than (1), the length of manual digit two

(Yates 2003a).

230. Ventrolateral twisting of the transverse axis of the

distal end of the first phalanx of manual digit one

relative to its proximal end: absent (0), present

but much less than 608 (1) or 608 (2) (Sereno

1999). Ordered.

231. Length of the first phalanx of manual digit one:

less than (0), or greater than (1), the length of the

first metacarpal (Gauthier 1986).

232. Shape of the proximal articular surface of the

first phalanx of manual digit one: rounded (0) or

with an embayment on the medial side (1)


233. Shape of the first phalanx of manual digit one:

elongate and subcylindrical (0) or strongly

proximodistally compressed and wedge shaped

(1) (Wilson 2002).

234. Length of the penultimate phalanx of manual

digit two: less than (0), or greater than (1), the

length of the second metacarpal (Rauhut 2003).

235. Length of the penultimate phalanx of manual

digit three: less than (0), or greater than (1), the

length of the third metacarpal (Rauhut 2003).

236. Shape of non-terminal phalanges of manual

digits two and three: longer than wide (0) or as

long as wide (1) (Yates 2003a).

237. Shape of the unguals of manual digits two and

three: straight (0), or strongly curved with tips

projecting well below flexor margin of proximal

articular surface (1) (Sereno et al. 1993).

238. Length of the ungual of manual digit two: greater

than the length of the ungual of manual digit one

(0), 75–100% of the ungual of manual digit one

(1), less than 75% of the ungual of manual digit

one (2) or the ungual of manual digit two is absent

(3) (modified from Gauthier 1986). Ordered.

239. Phalangeal formula of manual digits four and

five: greater than (0), or less than (1), 2-0,

respectively (Gauthier 1986).

240. Strongly convex dorsal margin of the ilium:

absent (0) or present (1) (Gauthier 1986).

241. Cranial extent of preacetabular process of ilium:

does not (0), or does (1), project further forward

than cranial end of the pubic peduncle (Yates

2003a).

242. Shape of the preacetabular process: blunt and

rectangular (0) or with a pointed, projecting

cranioventral corner and a rounded dorsum (1)


243. Depth of the preacetabular process of the ilium:

much less than (0), or subequal to (1), the depth

of the ilium above the acetabulum (modified

from Gauthier 1986).

244. Length of preacetabular process of the ilium: less

than (0), or greater than (1), twice its depth.

245. Buttress between preacetabular process and the

supra-acetabular crest of the ilum: present (0) or


246. Medial wall of acetabulum: fully closing acet-

abulum with a triangular ventral process

between the pubic and ischial peduncles (0),

partially open acetabulum with a straight ventral

margin between the peduncles (1), partially open

acetabulum with a concave ventral margin

between the peduncles (2) or fully open

acetabulum with medial ventral margin closely

approximating lateral rim of acetabulum (3)


247. Length of the pubic peduncle of the ilium: less

than (0), or greater than (1), twice the cranio-

caudal width of its distal end (Sereno 1999).


248. Caudally projecting ‘heel’ at the distal end of the

ischial peduncle: absent (0) or present (1) (Yates

2003b).

249. Length of the ischial peduncle of the ilium:

similar to pubic peduncle (0), much shorter than

pubic peduncle (1), or virtually absent so that

the chord connecting the distal end of the pubic

peduncle with the ischial articular surface

contacts the postacetabular process (Upchurch

et al. 2004). Ordered.

250. Length of the postacetabular process of the

ilium: between 40 and 100% of the distance

between the pubic and ischial peduncles (0), less

than 40% of this distance (1), or more than

100% of this distance.

251. Well developed brevis fossa with sharp margins

on the ventral surface of the postacetabular

process of the ilium: absent (0) or present (1)

(Gauthier 1986).

252. Anterior end of ventrolateral ridge bounding

brevis fossa: not connected to (0), or joining (1)

supracetabular crest (1).

253. Shape of the caudal margin of the postacetabular

process of the ilium: rounded to bluntly pointed

(0), square ended (1) or with a pointed ventral

corner and a rounded caudodorsal margin (2)

(Yates 2003b). Unordered.

254. Width of the conjoined pubes: less than (0), or

greater than (1), 75% of their length (Cooper,

1984).

255. Pubic tubercle on the lateral surface of the

proximal pubis: present (0) or absent (1) (Yates

2003a).

256. Proximal anterior profile of pubis: anterior margin

of pubic apron smoothly confluent with anterior

margin of iliac pedicel (0) iliac pedicel set anterior

to the pubic apron creating a prominent inflection

in the proximal anterior profile of the pubis (1).

257. Minimum transverse width of the pubic apron:

much more than (0), or less than (1), 40% of the

width across the iliac peduncles of the ilium.

258. Position of the obturator foramen of the pubis: at

least partially occluded by the iliac pedicel (0), or

completely visible (1), in anterior view (Galton

and Upchurch 2004).

259. Lateral margins of the pubic apron in anterior

view: straight (0) or concave (1) (Yates and

Kitching 2003).

260. Orientation of distal third of the blades of the

pubic apron: confluent with the proximal part of

the pubic apron (0) twisted posterolaterally

relative to proximal section so that the anterior

surface turns to face laterally (1) (Langer 2004).

261. Orientation of the entire blades of the pubic

apron: transverse (0) or twisted posteromedially


262. Craniocaudal expansion of the distal pubis:

absent (0), less than 15% (1), or greater than

15% (2), of the length of the pubis (modified

Gauthier 1986). Ordered.

263. Notch separating posteroventral end of the

ischial obturator plate from the ischial shaft:

present (0) or absent (1) (Rauhut 2003).

264. Elongate interischial fenestra: absent (0) or

present (1) (Yates 2003b).

265. Longitudinal dorsolateral sulcus on proximal

ischium: absent (0) or present (1) (Yates

2003a).

266. Shape of distal ischium: broad and plate-like, not

distinct from obturator region (0) or with a

discrete rod-like distal shaft (1).

267. Length of ischium: less than (0) or greater than

(1) that of the pubis (Salgado et al. 1997).

268. Ischial component of acetabular rim: larger than

(0) or equal to (1) pubic component (Galton and

Upchurch 2004).

269. Shape of the transverse section of the ischial

shaft: ovoid to subrectangular (0) or triangular

(1) (Sereno 1999).

270. Orientation of the long axes of the transverse

section of the distal ischia: meet at an angle (0) or

are coplanar (1) (Wilson and Sereno 1998).

271. Depth of the transverse section of the ischial

shaft: much less than (0) at least as great as (1),

the transverse width of the section (Wilson and

Sereno 1998).

272. Distal ischial expansion: absent (0) or present

(1) (Holtz 1994).

273. Transverse width of the conjoined distal ischial

expansions: greater than (0), or less than (1),

their sagittal depth (Yates 2003a).

274. Length of the hindlimb: greater than (0), or less

than (1), the length of the trunk (Gauthier

1986).

275. Longitudinal axis of the femur in lateral view:

strongly bent with an offset between the

proximal and distal axes greater than 158 (0),

weakly bent with an offset of less than 108 (1) or

straight (2) (Cooper, 1984). Ordered.

276. Shape of the cross section of the midshaft of the

femur: subcircular (0) or strongly elliptical with

the long axis oriented mediolaterally (1) (Wilson

and Sereno 1998).

277. Angle between the long axis of the femoral head

and the transverse axis of the distal femur: about

308 (0) or close to 08 (1) (Carrano, 2000).

278. Shape of femoral head: roughly rectangular in

profile with a sharp medial distal corner (0)

roughly hemispherical with no sharp medial

distal corner (1).

This character only applies to taxa with a

medially, or anteromedially protruding femoral

head. It does not apply to outgroup taxa

(Euparkeria or Crurotarsi) with proximally

directed femoral heads and is coded as unknown

in these taxa.

A. M. Yates22

279. Posterior proximal tubercle on femur: well-

developed (0) or indistinct to absent (1)

(Novas 1996).

280. Shape of the lesser trochanter: small rounded

tubercle (0), proximodistally oriented, elongate

ridge (1) or absent (2) (modified from Gauthier

1986). Unordered.

281. Position of proximal tip of lesser trochanter: level

with (0), or distal to (1) femoral head (Galton

and Upchurch 2004).

282. Projection of the lesser trochanter: just a scar

upon the femoral surface (0) or a raised

process (1).

283. Transverse ridge extending laterally from the

lesser trochanter: absent (0) or present (1)

(Rowe and Gauthier 1990).

284. Height of the lesser trochanter in cross section:

less than (0) or at least as high as basal width (1)

(modified from Galton 1990).

285. Position of the lesser trochanter: near the center

of the anterior face (0), or close to the lateral

margin (1), of the femoral shaft in anterior view.

286. Visibility of the lesser trochanter in posterior

view: not visible (0) or visible (1) (Galton and

Upchurch 2004).

287. Height of the fourth trochanter: tall crest (0) or a

low rugose ridge (1) (Gauthier 1986).

288. Position of the fourth trochanter along the length

of the femur: in the proximal half (0) or

straddling the midpoint (1) (Galton 1990).

289. Symmetry of the profile of the fourth trochanter

of the femur: sub symmetrical without a sharp

distal corner (0) or asymmetrical with a stee-

per distal slope than the proximal slope and a

distinct distal corner (1) (Langer 2004).

290. Shape of the profile of the fourth trochanter of

the femur: rounded (0) or subrectangular (1).

291. Postion of fourth trochanter along the medio-

lateral axis of the femur: centrally located (0) on

the medial margin (1) (Galton 1990).

292. Extensor depression on anterior surface of the

distal end of the femur: absent (0) or present (1)

(Molnar et al. 1990).

293. Size of the medial condyle of the distal femur:

subequal to (0) or larger than (1) the

fibularþ lateral condyles (modified from Wilson

2002).

294. Tibia: femur length ratio: greater than 1.0 (0),

between 1.0 and 0.6 (1) or less than 0.6 (2)


295. Orientation of cnemial crest: projects anteriorly

to anterolaterally (0) or projecting laterally (1)


296. Paramarginal ridge on lateral surface of cnemial

crest: absent (0) or present (1).

297. Position of the tallest point of the cnemial crest:

close to the proximal end of the crest (0) or about

half way along the length of the crest, creating an

anterodorsally sloping proximal margin of the

crest (1).

298. Proximal end of tibia with a flange of bone that

contacts the fibula: absent (0) or present (1)

(Gauthier 1986).

299. Position of the posterior end of the fibular

condyle on the proximal articular surface tibia:

anterior to (0) or level with posterior margin of

proximal articular surface.

300. Shape of the proximal articular surface of the

tibia: ovoid, anteroposteriorly longer than trans-

versely wide (0) or subcircular and as wide

transversely as anteroposteriorly long (Wilson

and Sereno 1998).

301. Transverse width of the distal tibia: subequal to

(0), or greater than (1), its craniocaudal length

(Gauthier 1986).

302. Anteroposterior width of the lateral side of the

distal articular surface of the tibia: as wide (0), or

narrower than (1), the anteroposterior width of

the medial side.

303. Relationship of the posterolateral process of the

distal end of the tibia with the fibula: not flaring

laterally and not making significant contact with

the fibula (0), flaring laterally and backing the

fibula (1).

304. Shape of the distal articular end of the tibia in

distal view: ovoid (0) or subrectangular (1).

305. Shape of the anteromedial corner of the distal

articular surface of the tibia: forming a right

angle (0) or forming an acute angle (1) (Langer

2004).

306. Position of the ateral margin of descending

caudoventral process of the distal end of the

tibia: protrudes laterally at least as far as (0), or

set well back from (1), the craniolateral corner of

the distal tibia (Wilson and Sereno 1998).

307. A triangular rugose area on the medial side of the

fibula: absent (0) or present (1) (Wilson and

Sereno 1998).

308. Transverse width of the midshaft of the fibula:

greater than 0.75 (0), between 0.75 and 0.5 (1),

or less than 0.5 (2), of the transverse width of

the midshaft of the tibia (Langer 2004).

Ordered.

309. Position of fibula trochanter: on anterior surface

of fibula (0), laterally facing (1), or anteriorly

facing but with strong lateral bulge (2) (modified

from Wilson and Sereno 1998).

310. Depth of the medial end of the astragalar body in

cranial view: roughly equal to the lateral end (0)

or much shallower creating a wedge shaped

astragalar body (Wilson and Sereno 1998).

311. Shape of the caudomedial margin of the

astragalus in dorsal view: forming a moderately

sharp corner of a subrectangular astragalus (0) or

evenly rounded without formation of a caudo-

medial corner (1) (Wilson and Sereno 1998).


312. Dorsally facing horizontal shelf forming part of

the fibular facet of the astragalus: present (0) or

absent with a largely vertical fibular facet (1)

(Sereno 1999).

313. Pyramidal dorsal process on the posteromedial

corner of the astragalus: absent (0) or present (1).

314. Shape of the ascending process of the astragalus:

anteroposteriorly deeper than transversely wide

(0) or transversely wider than anteroposteriorly

deep (1).

315. Posterior extent of ascending process of the

astragalus: well anterior to (0), or close to

posterior margin (1), of astragalus (Wilson and

Sereno 1998).

316. Sharp medial margin around the depression

posterior to the ascending process of the

astragalus: absent (0) or present (1) (Novas

1996).

317. Buttress dividing posterior fossa of astragalus

and supporting ascending process: absent (0) or

present (1) (Wilson and Sereno 1998).

318. Vascular foramina set in a fossa at the base of the

ascending process of the astragalus: present (0)

or absent (1) (Wilson and Sereno 1998).

319. Transverse width of the calcaneum: greater

than (0), or less than (1), 30% of the transverse

width of the astragalus (Yates and Kitching

2003).

320. Lateral surface of calcaneum: simple (0) or with

a fossa (1).

321. Medial peg of calcaneum fitting into astragalus:

present, even if rudimentary (0) or absent (1)

(Sereno et al. 1993).

322. Calcaneal tuber: large and well-developed (0) or

highly reduced to absent (1).

323. Shape of posteromedial heel of distal tarsal four

(lateral distal tarsal): proximodistally deepest

part of the bone (0) or no deeper than the rest of

the bone (1) (Sereno et al. 1993).

324. Shape of posteromedial process of distal tarsal

four in proximal view: rounded (0) or pointed

(1) (Langer 2004).

325. Ossified distal tarsals: present (0) or absent (1)

(Gauthier 1986).

326. Proximal width of the first metatarsal: is less than

(0), or at least as great as (1), the proximal width

of the second metatarsal (modified from Wilson

and Sereno 1998).

327. Orientation of proximal articular surface of

metatarsal one: horizontal (0), or sloping

proximolaterally relative to the long axis of the

bone (1) (Wilson 2002).

328. Orientation of the transverse axis of the distal

end of metatarsal one: horizontal (0), or angled

proximomedially (1) (Wilson 2002).

329. Shape of the medial margin of the proximal

surface of the second metatarsal: straight (0) or

concave (1) (modified from Sereno 1999).

330. Shape of the lateral margin of the proximal

surface of the second metatarsal: straight (0) or

concave (1) (modified from Sereno 1999).

331. Length of the third metatarsal: greater than (0),

or less than (1), 40% of the length of the tibia

(Gauthier 1986).

332. Minimum transverse shaft diameters of third and

fourth metatarsals: greater than (0), or less than

(1), 60% of the minimum tansverse shaft

diameter of the second metatarsal (Wilson and

Sereno 1998).

333. Transverse width of the proximal end of the

fourth metatarsal: less than (0), or at least (1),

twice the anteroposterior depth of the proximal

end (modified from Sereno 1999).

334. Transverse width of the proximal end of the fifth

metatarsal: less than 25% (0), between 30 and

49% (1), or greater than 50% (2), of the length

of the fifth metatarsal (modified from Sereno

1999). Ordered.

335. Transverse width of distal articular surface

of metatarsal IV in distal view: greater (0), or

less (1), than anteroposterior depth (Sereno

1999).

336. Pedal digit five: reduced, non-weight bearing (0),

or large (fifth metatarsal at least 70% of fourth

metatarsal), robust and weight bearing (1)


337. Length of non-terminal pedal phalanges: all

longer than wide (0), proximal-most phalanges

longer than wide while more distal phalanges are

as wide as long (1) or all non-terminal phalanges

are as wide, if not wider, than long (2) (modified

from Wilson and Sereno 1998). Ordered.

338. Length of the first phalanx of pedal digit one:

greater than (0), or less than (1) the length of the

ungual of pedal digit one (Yates and Kitching

2003).

339. Length of the ungual of pedal digit one: less than

at least some non-terminal phalanges (0) or

longer than all non-terminal phalanges (1).

340. Shape of the ungual of pedal digit one: shallow,

pointed, with convex sides and a broad ventral

surface (0), or deep, abruptly tapering, with

flattened sides and a narrow ventral surface (1)


341. Shape of proximal articular surface of pedal

unguals: proximally facing, visible on medial and

lateral sides (0) or proximomedially facing and

visible only in medial view, causing medial

deflection of pedal unguals in articulation (1)


342. Penultimate pahalanges of pedal digits two and

three: well developed (0) or reduced disc-shaped

elements if they are ossified at all (1) (Wilson and

Sereno 1998).

343. Shape of the unguals of pedal digits two and

three: dorsoventrally deep with a proximal

A. M. Yates24

articulating surface that is at least as deep as it is

wide (0) or dorsoventrally flattened with a

proximal articulating surface that is wider than

deep (1) (Wilson and Sereno 1998).

344. Length of the ungual of pedal digit two: greater

than (0), between 100% and 90% (1), or less

than 90% (2), of the length of the ungual of pedal

digit one (modified from Gauthier 1986).

Ordered.

345. Size of the ungual of pedal digit three: greater

than (0), or less than (1), 85% of the ungual of

pedal digit two in all linear dimensions (Yates

2003a).

346. Number of phalanges in pedal digit four: four (0)

or fewer than four (1) (Gauthier 1986).

347. Phalanges of pedal digit five: present (0) or


348. Femoral length: less than 200 mm (0), between

200 and 399 mm (1), between 400 and 599 mm

(2), between 600 and 799 mm (3), between 800

and 1000 mm (4) or greater than 1000 mm

(modified from Yates 2004).

References

Benton MJ, Juul L, Storrs GW, Galton PM. 2000. Anatomy and

systematics of the prosauropod dinosaur Thecodontosaurus

antiquus from the upper Triassic of southwest England. J Vert

Paleontol 20:77–108.

Bonaparte JF. 1986. The early radiation and phylogenetic

relationships of the Jurassic sauropod dinosaurs, based on

vertebral anatomy. In: Padian K, editor. The beginning of the

age of dinosaurs. Cambridge: Cambridge University Press.

p 247–258.

Galton PM. 1985. Cranial anatomy of the prosauropod dinosaur

Sellosaurus gracilis from the middle Stubensandstein (upper

Triassic) of Nordwurttemberg, West Germany. Stuttgart

Beitrage zur Naturkunde (B) 118:1–39.

Galton PM. 1990. Basal sauropodomorpha—prosauropods. In:

Weishampel DB, Dodson P, Osmolska H, editors. The

dinosauria. Berkeley: University of California Press.

p 320–344.

Galton PM, Upchurch P. 2004. Prosauropoda. In: Weishampel DB,

Dodson P, Osmolska H, editors. The Dinosauria. 2nd ed.


Gauthier J. 1986. Saurischian monophyly and the origin of birds.

Memoirs Californian Acad Sci 8:1–55.

Holtz Jr TR. 1994. The phylogenetic position of Tyrannosauridae:

Implications for theropod systematics. J Paleontol

68:1100–1117.

Langer MC. 2004. Basal Saurischia. In: Weishampel DB, Dodson P,

Osmolska H, editors. The Dinosauria. 2nd ed. Berkeley:

University of California Press. p 25–46.

Leal LA, Azevedo SAK, Kellner AWA, Da Rosa AAS. 2004. A

new early dinosaur (Sauropodomorpha) from the Caturrita

Formation (Late Triassic), Parana Basin, Brazil. Zootaxa

690:1–24.

Molnar RE, Kurzanov SM, Dong Z. 1990. Carnosauria.

In: Weishampel DB, Dodson P, Osmolska H, editors.

The Dinosauria. Berkeley: University of California Press.

p 169–209.

Novas FE. 1993. New information on the systematics and

postcranial skeleton of Herrerasaurus ischigualastensis

(Theropoda: Herrerasauridae) from the Ischigualasto

Formation (Upper Triassic) of Argentina. J Vert Paleontol

13:400–423.

Novas FE. 1996. Dinosaur monophyly. J Vert Paleontol

16:723–741.

Perez-Moreno BP, Sanz JL, Buscalioni AD, Moratalla JJ, Ortega F,

Rasskin-Gutman D. 1994. A unique multitoothed ornithomi-

mosaur dinosaur from the Lower Cretaceous of Spain. Nature

370:363–367.

Rauhut OWM. 2003. The interrelationships and evolution of basal

theropod dinosaurs. Special papers in palaeontology 69.

London: The Palaeontological Association.

Sereno PC. 1999. The evolution of dinosaurs. Science

284:2137–2147.

Sereno PC, Forster CA, Rogers RR, Monetta AM. 1993. Primitive

dinosaur skeleton from Argentina and the early evolution of

dinosauria. Nature 361:64–66.

Sereno PC, Dutheil DB, Iarochene M, Larsson HCE, Lyon GH,

Magwene PM, Sidor CA, Varricchio DJ, Wilson JA. 1996.

Predatory dinosaurs from the sahara and Late Cretaceous faunal

differentiation. Science 272:986–991.

Upchurch P. 1995. The evolutionary history of sauropod dinosaurs.

Philos Trans R Soc Lond B 349:365–390.

Upchurch P. 1998. The phylogenetic relationships of sauropod

dinosaurs. Zool J Linnean Soc 124:43–103.

Upchurch P, Barrett PM, Dodson P. 2004. Sauropoda. In:

Weishampel DB, Dodson P, Osmolska H, editors. The

Dinosauria. 2nd ed. Berkeley: University of California Press.

p 259–322.

Wilson JA. 2002. Sauropod dinosaur phylogeny: critique and

cladistic analysis. Zool J Linnean Soc 136:217–276.

Wilson JA, Sereno PC. 1998. Early evolution and higher-level

phylogeny of sauropod dinosaurs. Soc Vert Paleontol Memoirs

5:1–68.

Yates AM. 2003a. A new species of the primitive dinosaur,

Thecodontosaurus (Saurischia: sauropodomorpha) and its impli-

cations for the systematics of early dinosaurs. J Syst Palaeontol

1:1–42.

Yates AM. 2003b. The species taxonomy of the sauropodomorph

Dinosaurs from the Lowenstein Formation (Norian, Late

Triassic) of Germany. Palaeontology 46:317–337.

Yates AM. 2003c. The first definite prosauropod dinosaur from the

Lower Elliot Formation (Norian: Upper Triassic), South Africa.

Palaeontologia Africana 39:63–68.

Yates AM. 2004. Anchisaurus polyzelus (Hitchcock): the smallest

known sauropod dinosaur and the evolution of gigantism

amongst sauropodomorph dinosaurs. Postilla 230:1–58.

Yates AM, Kitching JW. 2003. The earliest known sauropod

dinosaur and the first steps towards sauropod locomotion. Proc

R Soc Lond B 270:1753–1758.

Character-taxon matrix

Euparkeria

0 0 0 0 0 0 0 0 ? 0 0 0 ? 0 0 0 0 0 0 0 ? 0 0 0 ? 1 0 0 1 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 ? 0 0 0 ? 0 0 0 0

0 0 0 0 0 ? 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0

0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? 0 0 0 0 0 0 0 0 0 ? 0 0 ? 0 ? 0 0 ?

? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 ? 0 ? ? ? ?

? ? ? ? ? 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? 0 0 0 0 0 0

0 0 0 0 1 ? 0 ? 0 ? 0 0 0 0 ? ? 0 0 ? 0 0 0 0 ? 0 0 0 0 ? 0 0 1

0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 ? 0 2 ? 0 0 0 ? ? 0

1 0 0 0 0 0 1 ? 0 0 0 ? 1 0 0 0 0 0 ? 0 0 0 0 0 0 0 ? ? 0 ? 0 0

0 1 0 ? ? 0 0 0 0 ? ? 0 0 0 1 ? 0 0 0 0 0 0 0 0 0 ? 0 0 0


Crurotarsi

0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0&1 0 0 0 0 0 0 0 0 0 ? 0 0 0 0

0 0 0 0 0 0&1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0 0 0 0 ? ? 0 0 0 0 0 2 0 0 0 ? 0 0

? 0 ? 0 0 ? ? 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0&1 0 0 ? 0 0&1 0 0 0 2 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0&1 0 0

0 0 0 0 0 0 ? 0 0 0 0 ? 0 0&1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 ? 0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 1 ? 0 0 0 0 0

0 0 0 0 0 ? 0 0 0 0 0 0 0 ? ? 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0&1

Marasuchus

0 0 ? ? ? ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? 0 0 ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 ? ?

0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 ? 0 0 0 0 0

0 0 0 ? 1 ? 1 0 0 0 0 0 0 0 0 0 1 0 ? ? ? 0 0 ? 0 ? ? 0 0 ? 0 0 0

0 0 0 ? 0 0 ? ? ? 0 0 ? ? 0 0 0 ? 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 0

0 0 0 ? ? 0 1 0 1 0 0 0 0 0 0 0 ? 0 0 0 0 1 2 1 0 0 0 ? 0 0 0 0

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 0 0 0 1 0 0

0 0 0 0 ? 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0

0 0 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0

0 1 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 1 0 ? 0 ? ?

0 0 ? ? ? 1 0

Agnosphitys

? 0 ? ? ? ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 ? 0 ? 0 0 0 1 0 0 ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 ? 0 0 0 0 0

0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? 1 1 0 0 ? 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? 0 0 0 0 0 1 2 0 0 0 0 1 0 0 ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 0

? ? ? ? ? ? 0 1 1 0 0 0 1 0 0 ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ?

Anchisaurus

1 0 ? ? ? 0 0 ? ? 0 1 2 ? 1 ? ? 1 1 ? ? ? ? ? 1 1 1 0 1 1 ? 0 1 0 ?

0 1 0 1 0 1 1 0 0 0 1 1 0 0 1 1 1 1 0 1 2 0 0 ? 1 0 ? ? ? ? ? 2

0 1 ? 0 0 1 1 0 1 1 ? 2 0 0 2 0 1 0 ? ? ? ? ? ? 1 0 ? 1 0 ? 0 0 ?

? 1 0 0 ? 0 0 1 0 1 0 1 ? 1 1&2 ? 0 1 1 1 ? 0 0 1 1 ? ? ? 1 0 0

1 0 0 1 1 1 1 0 1 1 ? 1 0 0 0 0 ? 0 1 0 ? 0 0 1 1 0 0 ? 0 0 0 ?

? 0 0 0 0 0 0 0 ? ? 0 1 0 0 ? 0 ? ? 0 ? 0 1 ? 0 0 ? 1 0 ? ? ? ? ? 1

1 0&1 ? ? 2 1 1 0 0 1 0 1 1 0 0 ? ? 0 ? ? 1 0 0 1 0 1 1 1 0 0

1 0 0 2 0 1 0 0 0 0 0 2 0 0 0&1 1 0 1 0 3 1 0 0 0 0 0 ? 1 1

1 1 1 0 0 0 1 0 0 1 1 ? 0 0 1 0 1 0 0 0 0 1 0 1 1 0 1 0 0 0 0

1 0 0/1 1 1 1 0 1 0 0 0 0 0 0 1 1 0 1 1 1 ? 1 ? 0 1 ? 0 1 0 ? ?

? 1 0 1 1 1 ? 0 ? 0 1 ? ? 0 0 1 1 ? 1 0 1 0 0 0 0 0 1 0 0 1 1

Antetonitrus

? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 ? 1 0

1 2 0 0 0 ? ? ? ? ? ? ? ? 0 ? 0&1 1 ? ? ? ? ? ? ? ? 0 ? ? 0 0 ? 0

0 0 0 0 1 ? ? 0 0 2 1 0 0 1 0 0 1 0 1 2 1 0 0 1 0 0 0 ? ? ? ?

? ? ? ? 1 0 0 0 0 ? 0 1 1 0 0 ? ? ? 0 ? 0 0 1 ? ? 3 1 1 0 0 0 1

0 1 1 0 ? ? ? ? ? ? 0 0 1 0 ? 3 1 0 0 ? ? ? 1 0 1 0 ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 1 0 1 0 0 0 1 ? ? ? ? ? ? ? ? ? ? ? ? 1

1 1 0 1 1 1 1 0 0 1 1 1 1 1 1 1 1 0 1 0 0 0 0 0 0 1 1 0 1 1

1 0 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 0 1 1 1 1 0 ? ? ? ? 1&2

1 ? 0 0 ? 0 ? ? ? ? 3

Barapasaurus

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 ? 1 1 1 2 1 1 ? ?

? ? ? ? ? ? ? ? ? 1 ? 1 ? ? ? ? 0 0 ? ? 0 0 2 ? ? 1 0 0 1 0 1 ? ? 0 0

2 0 1 1 0 1 1 1 1 1 2 0 1 2 1 1 0 0 ? 0 1&2 1 0 0 1 ? 1 0 ? 1

1 ? ? 1 ? ? ? ? ? ? ? ? 1 0 1 0 0 ? 1 ? ? ? ? ? ? ? 1 1 ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 1 1 0 0 3 1 0 1 1 0 0 0 0 1

1 1 1 0 0 1 1 ? ? ? 1 1 1 1 0 0 1 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? 1 ? 1 ? 1 ? ? ? ? ? ? ? ? ? 1 ? 1 ? ? ? ? ? ? ? 1 ? ? ? ? ? ? ? ? ? 0 ?

? ? ? ? ? ? ? ? ? ? 1 ? 1 ? ? ? ? ? ? 5

Blikanasaurus

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 0 0 ? 0 1 1 0 1 1 1 ? 1 ? 0 0 0 0 1

0 1 1 0 1 0 1 1 1 0 0 1 0 1 1 1 1 1 1 2 0 1 2 1 1 0 0 0 0 1 0

0 ? 2

Camelotia

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? 1 ? ? ? ? ? ? ? ? ? ? ? 0 ? ? 0 ? ? 0 0 1 ? ? 0 0 2 1 0

0 0 0 0 0 0 0 ? ? 0 0 ? 0 ? ? ? ? ? ? ? ? ? ? 1 ? ? ? 0 ? ? 0 1 1 ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 ? ? ? ? ? ? ? ? ? ? 1 ? ? 0

0 1 1 1 ? 1 1 1 0 1 1 1 1 0 1 1 1 ? 1 ? ? 1 1 0 ? 0 0 0 0 0 0 ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1&2 ?

? ? 0 0 0 ? ? ? ? 5

A. M. Yates26

Cetiosaurus

1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

2 ? ? 0 0 1 1 1 1 0 1 1 1 1 1 0 1 0 0 1 0 0 0 2 0 ? 2 0 0 1 ? 1

0 ? 1 0 2 0 ? ? 1 0 1 1 1 0 2 0 1 0 1 0 0 ? ? ? 1&2 ? ? ? ? ? 1

0 0 1 ? ? 0 0 1 0 0 ? ? ? 0 1 1 1 1 1 0 3 1 1 0 0 1 0 0 0 1 1 ?

? ? ? ? ? ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 1 1 0 0 3 ? 0 2

1 0 ? 0 0 1 0 ? 1 ? 0 1 1 ? ? 1 1 0 1 ? 0 0 ? ? ? 2 1 1 1 1 2 ? ?

0 ? ? ? 0 1 0 ? 1 1 1 2 1 0 1 0 0 0 1 0 0 1 0 1 1 1 ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 5

Chindesaurus

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? 1 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 ? 0 ? ? 0 ? 1 ? ? 0 ? 1 ? ? ?

? ? ? 0 0 0 ? ? 0 0 ? ? ? ? ? ? ? ? ? ? ? 1 ? ? 0 ? ? ? ? 1 ? 0 0 ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? ? 1 0 1 ? 1 ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ?

? ? 0 0 ? 0 1 0 1 1 1 0 0 0 0 0 0 0 1 0 0 ? ? 0 0 ? 1 0 1 1 1 1

0 0 ? ? ? 0 1 1 1 1 0 1 0 0 ? ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? 1

Coloradisaurus

? 0 0 ? 1 0 0 2 ? 0 1 2 ? 1 1 1 1 1 1 ? 1 ? ? 1 1 1 0 ? 1 1 1 1

0 0 ? 1 0 ? ? 0 0 ? 0 0 1 ? 0 1 1 1 0 1 0 1 ? 1 0 0 1 0 0 1 0 0

? 1 0 1 1 0 1 1 0 1 1 ? ? 1 1 0 1 1 1 ? ? ? 0 ? 1 ? 1 0 0 1 0 1

1 0 1 0 1 1 1 0 2 1 0 0 1 0 1 0 1 0 0 0 0 ? 1 ? 0 0 ? 1 1 0 0

2 0 0 1 0 0 0 1 1 ? 0 1 1 0 ? ? 0 0 ? 0 0 1 ? 1 0 0 1 1 0 0 0 0

0 0 0 0 0 0 0 0 0 0 1 0 1 1 1 0 ? ? 0 1 0 0 1 0 0 ? 0 ? ? ? ? ? ?

? ? ? 0 1 1 1 1 1 1 2 1 ? 0 1 1 ? ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? 0 0 1 0 0 0 3 1 1 0 0 ? 0 ? 1 1 1 0 1 1 0 0 1

0 0 1 1 1 1 1 0 1 1 1 ? 0 0 1 0 1 1 0 1 0 0 0 0 1 0 1 1 1 1 0

1 0 0 0 0 ? 0 1 1 0 1 1 0 ? ? ? 0 1 1 1 1 0 ? 0 0 0 1 0 1 ? ? ?

0 0 1 1 1 0 0 1 2 ? 1 0 1 ? 0 0 0 0 2 1 0 0 2

Efraasia

1 0 0 ? 1 0 0 1 ? 0 1 ? ? 1 ? 1 1 1 ? 1 1 2 ? 1 1 1 0 ? 1 0 0 ? ?

? 0 1 0 0 1 0 0 0 0 0 ? ? 0 ? 1 0 ? ? 0 1 ? 1 ? ? 1 ? ? ? ? ? ? 1 0

? 0 0 0 ? 1 ? ? ? ? 0 1 1 0 1 0 0 ? ? ? ? ? ? 1 0 0 1 0 0 1 0 ? ? 1

1 ? 0 2 1 0 0 1 0 1 0 1 0 0 0 0 1 ? ? ? 1 ? 1 1 0 0 1 1 0 1 0 0

1 0 1 1 0 1 1 0 1 0 0 0 0 0 0 1 ? ? 0 0 1 ? 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 0 1 0 0 0 ? 0 1 1 0 0 0 0 ? 0 ?

0 1 0 1 1 2 1 2 0 0 0 1 1 1 0 0 1 1 0 1 0 1 2 0 1 0 ? 1 1 0 0

0 0 0 1 1 1 0 1 0 0 0 1 0 0 0 1 0 0 0 3 1 0 0 0 1 0 1 1 0 0 0

1 0 0 0 0 0 0 1 1 0 0 1 0 1 1 0 1 0 0 0 0 1 1 1 1 0 0 0 0 1 0

1 1 0 0 0 1 0 0 0 0 0 0 1 1 0 1 ? 0 0 1 ? 0 1 1 0 1 0 ? 0 ? 0 1

? 1 1 1 0 0 0 1 ? ? 0 0 1 1 ? 1 0 1 0 0 0 0 0 1 0 0 0 2

Eoraptor

0 0 1 0 ? 0 0 0 ? 0 1 1 0 ? 0 0 1 1 0 0 0 0 0 1 0 1 0 0 1 0 0

1 ? 0 0 1 1 0 1 0 0 0 0 0 ? 1 0 1 1 0 0 1 0 1 0 0 0 0 1 0 0 0

0 0 1 0 0 1 ? ? ? 0 ? ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 0 0 0 0 ?

0 ? 0 ? 0 ? 0 ? 0 0 ? 1 0 0 0 0 0 0 0 0 ? ? ? ? ? ? 1 0 0 0 0 1 0

1 0 0 0 0 ? 1 0 ? ? ? ? 0 0 0 ? ? 0 0 ? 0 ? ? ? ? 0 0 ? 0 0 0 ? ? 0

0 0 0 0 0 0 1 ? ? 1 0 ? ? ? 1 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 ? ? ?

0 1 1 ? ? ? 0 ? 0 ? ? ? ? ? ? ? ? 2 0 ? 0 0 0 ? 1 ? 0 0 0 ? 0 ? 0 1

0 0 0 0 1 1 0 1 0 0 ? 2 ? 0 0 1 1 1 0 ? ? 0 ? ? ? 0 ? 0 ? ? ? 1 0

? 0 0 1 0 ? 1 0 ? ? ? ? 0 ? 1 0 0 ? ? 0 0 1 1 ? ? 0 0 0 ? 0 ? 1 0 ?

0 1 1 0 ? ? 1 ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? 0 ? 0 ? ? ? 0 0 ? 0 0 1 0

0 0 0 0 0 0 ? ? 0 0 0

Eucnemesaurus

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? ? 0 1 ? ? 0 0 1 1 0 0

1 0 0 0 0 0 ? ? 0 0 ? 0 ? ? ? ? ? ? ? ? ? ? 1 0 0 0 0 ? 0 1 1 0 0 ?

? ? ? ? ? ? ? 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 0 ? ? ? 0 ? ? ? ? ? ? ?

? ? ? ? ? ? 0 0 1 1 0 1 ? 1 0 1 0 ? 1 0 1 0 1 1 0 ? ? 0 0 0 0 0 1

1 0 1 1 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? 3&4

Gongxianosaurus

1 ? ? ? ? ? 0 ? ? ? ? ? ? ? ? ? 1 2 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 ? ? ? 1 ? ? 2 1 2 1 ?

? ? ? ? ? ? ? ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? 1 ? ? 0 ? ? 0 0 1 ? ? 0 ? ?

? ? ? ? 0 ? 0 ? ? 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? 0 ? 0 ? 1 ? 0

? 0 0 0 ? 0 1 1 ? ? 2 1 1 0 ? ? ? ? 1 ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? 1 0 0 ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? 1&2 1 1 1 ? 2 ? ? ? ? ? ? 0 ? ? ? 1 1 0 1 ? ? ? ?

? ? 1 ? 0 ? ? 1 ? ? ? 0 ? 1 ? 1 ? ? ? ? 1 ? 1 1 ? ? 0 1 0 ? ? ? 1 0 ?

2 ? 0 1 1 1 0 0 0 0 2 0 0 0 5

Plateosaurus ( 5 Gresslyosaurus) ingens

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 ? ? 0 1 ? ? ? ? ? ? ? ? 1 0 ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 ? 0 1 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0&1 ? ? ? 0 ? 0 ? ? ? ? 4


Guaibasaurus

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? ? 0 1 ? ? 0 1 1 0 0 1

0 0 0 0 0 0 ? ? 0 0 0 0 0 ? ? ? ? ? ? ? 0 ? 0 1 1 0 0 ? ? ? ? ? ? ?

1 ? ? ? ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? 0 ? ? ? ? ? 1 ? 0 0 2 1 0 1 1 0 0 1 0 0 0 0 0 0 0

0 1 0 ? 1 0 1 1 0 ? 0 0 0 0 1 0 0 1 0 0 0 0 1 0 ? 1 ? 1 0 1 0 0

0 0 ? 0 0 1 1 1 1 0 0 1 ? 0 ? 0 1 ? ? ? ? 0 0 1 0 1 ? ? 0 0 0 0 ?

? 1 0 ? 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1

Herrerasaurus

0 0 0 0 0 0 0 0 ? 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 1 0 0

0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 1 0 1 0 1 0 1 0 0 0 1 0 0

0 0 1 1 0 1 0 0 0 0 0 0 1 1 ? 0 0 0 0 0 0 0 ? 1 0 0 0 ? 0 0 0

0 0 0 0 0 ? 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1

0 0 0 0 0 0 1 0 0 0 0 0 ? 0 0 0 0 1 0 0 0 0 0 1 1 ? ? 0 0 1 0

0&1 1 0 0 0 0 0 0 2 0 0 0 0 0 0 1 0 0 0 0 0 1 0 1 1 0 0 0 0

0 1 1 0 0 1 0 1 0 0 ? 1 1 1 ? ? 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0

0 1 2 0 0 0 0 0 1 1 0 0 0 0 0 1 1 0 1 1 0 1 0 1 0 0 0 0 0 1 2

0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 ? ? ? 0 1 ? 0 1 0 1 0 0 1 0 0 0 0

1 0 1 1 1 0 0 0 1 0 1 1 0 0 0 1 0 0 0 0 1 0 0 0 0 1 0 0 0 1 0

0 1 0 0 1 0 1 0 0 0 0 0 0 1 1 0 0 0 1 0 0 0 0 1 0 0 1 0 0 0 0

0 0 0 0 0 ? 0 2

Isanosaurus

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? 0 ? ? 0 ? ? ? ? ? ? ? ? ? ? ? 2 ? ? 1 0 ? ? 0 ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? 2 ? 0 1 1 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? 1 0 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? 2 1 1 1 1 2 ? ? 0 ? ? ? 0 0 1 ? 1 ? 0 ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? 3

Jingshanosaurus

1 0 0 1 ? 0 0 2 ? ? 1 2 1 1 1 1 1 1 0 0 ? 0 ? 1 1 1 0 1 1 1 0 1

1 0 ? 1 0 2 0 0 1 ? ? 1 ? 1 0 1 1 1 0 1 1 1 0 1 0 0 1 0 0 0 ? ?

? ? 0 1 1 0 0 0 0 0 1 1 ? 2 1 1 0 0 1 0 0 0 ? ? 1 ? 1 0 1 1 0 0

1 0 ? ? 1 1 1 0 2 1 0 0 1 0 1 2 1 1 0 0 1 ? ? ? ? 1 1 1 ? ? 0 1

0 ? ? ? ? ? ? 1 ? ? ? 1 0 1 0 0 ? ? 0 0 1 1 1 0 0 ? 1 ? ? ? 0 0 0 0

0 0 0 0 0 0 0 0 0 ? ? 1 0 ? ? 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0

? 0 0 1 ? ? 1 1 2 0 1 0 1 1 1 0 0 ? ? 0 0 ? ? 0 0 1 1 ? 3 1 0 0

1 0 0 ? 0 1 0 0 0 1 0 2 0 0 0 1 0 0 0 3 1 0 0 0 0 0 2 1 1 1 0

0 1 0 0 2 ? ? 1 1 0 0 1 0 1 1 0 ? 0 1 0 0 1 1 0 1 0 0 0 0 1 0

1 1 1 1 0 1 0 1 0 0 0 0 1 1 0 1 1 0 ? 1 2 0 1 1 0 1 0 ? 0 0 ? ?

? ? ? ? 0 1 0 1 1 1 1 0 1 2 0 1 0 1 0 0 0 0 0 0 1 0 ? 4

Lufengosaurus

1 0 0 ? ? ? 0 2 ? ? ? 2 1 1 ? 1 ? 1 1 ? 1 1 1 1 1 1 1 1 1 1 0 1 0

0 0 1 0 1 1 0 1 0 0 0 0 1 0 1 1 1 0 1 0 1 1 1 0 0 1 0 ? 1 ? ? ?

? 0 1 1 0 ? ? 0 1 1 ? ? 0 1 0 1 0 1 0 0 ? 0 1 1 0 1 0 0 1 0 ? ? ?

? 0 1 1 1 ? 2 1 0 1 1 0 1 0 1 0 0 0 1 1 ? 0 0 0 1 1 1 0 0 2 0

0 1 1 0 1 1 1 ? 0 1 1 0 1 0 0 0 2 0 0 1 1 1 0 0 1 1 0 0 0 0 0

0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 1 0 0 1 0 0 0 0 0 0 0 0&1 1 0

0 0 0 0 0 0 0 0 1 1 1 1 1 2 0&1 1 0 1 1 1 0 0 1 1 0 1 1 1 1

0 1 1 1 3 1 0 0 1 0 1 2 0 1 0 0 0 1 0 2 0 0 0 1 0 0 0 3 1 1 0

0 0 0 2 1 1 1 0 1 1 0 0 2 1 0 1 1 0 0 1 0 1 1 0 0 0 0 1 0 1 1

0 1 0 0 0 0 1 0 2 1 1 1 0 1 0 1 0 0 0 0 1 1 0 1 1 0 0 1 0 0 1

1 1 1 0 ? 0 0 0 1 ? 1 1 1 0 1 0 1 1 1 0 0 1 2 0 1 0 1 1 0 0 0

0 1&2 0&1 0 0 4

Mamenchisaurus

1 1 0 0 0 1 1 3 ? 1 1 2 0 1 1 0 1 2 0 0 0 0 0 0 0 1 0 1 1 ? 2

? ? ? 1 1 1 2 0 1 0 1 0 1 0 1 1 1 ? 1 ? 1 0 1 2 0 1 0 0 0 0 0 1

1 1 2 2 1 0 ? ? 0 ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 0&1 0 0 0

0 1 ? 0 1 0 0 0 1 1 1 1 1 1 1 1 1 2 1 1 1 3 ? ? 0 0 1 1 1 0 1

2 1 0 ? ? ? ? 0 1 1 1 0 0 2 ? 1 1 1 0 1 0 1 0 ? 0 0 2 0 ? 1 ? 0

0 1 ? 0 2 0 1 2 1 0 0 0 ? 0 1 1 ? ? 1 ? 1 1 1 1 1 1 0 1 ? 0 0 1

0 1 1 ? 1 ? ? ? ? ? ? ? 0 0 ? ? 0 ? ? ? ? ? ? ? ? ? ? 1 ? ? ? ? ? ? ? ? ?

? ? ? ? 1 ? ? ? ? ? ? 1 1 1 1 0 0 3 1 0 1 1 ? 0 0 ? 1 1 ? 1 0 0 1 1

0 0 ? 1 1 ? ? 0 0 1 0 0 2 1 1 1 1 2 ? ? 0 ? ? ? 0 1 0 ? 1 1 1 2 1

0 1 0 ? 1 ? ? 0 ? ? ? ? ? ? 1 0 1 0 1 1 1 1 1 ? ? ? ? ? ? 1 ? ? ? ? ?

2 ? ? 2 ? 0 2 ? 1 1 1 1 0 2 ? ? 0 5

Massospondylus

1 0 0 1 1 0 0 2 1 0 1 2 1 1 1 1 1 1 1 1 1 2 1 1 0 1 1 1 1 1 0

1 0 0 0 1 0 1 1 0 1 0 0 0 1 1 0 0 1 1 0 1 1 1 1 1 0 0 1 0 0 0

1 0 2 ? 0 1 ? 0 1 1 0 0 1 1 1 0 1 0 1 0 1 ? ? 1 0 1 0 0 1 0 0 1

0 1 1 0 1 0 1 0 1 0 2 1 0 0&1 1 0 1 0 1 1 0 0 1 1 1 0 0 ? 1

1 1 0 0 2 0 0 1 0 0 1 1 1 1 0 1 1 0 1 0 0 0 2 0 0 1 1 1 0 0 1

1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 1 0 1 0 0 1 0&1 0 0 0

0 0 0 0 0 0 0 0 0 0 0 ? 0 1 0&1 1 1 1 1 2 0 1 0 1 1 1 0 0 1

1 0 1 1 1 1 0 1 1 1 2 1 0 0 1 0 1 2 1 1 0 0 0 0 0 2 0 0 0 1 0

0 0 3 1 0 0 0 1 0 0 1 1 1 0 1 1 0 0 1 0 0 1 1 0 0 1 0 1 1 1 0

0 0 1 0 1 1 0 1 0 0 0 0 1 0 2 1 0 1 0 1 0 0 0 0 0 0 1 1 0 1 1

0 0 1 0 0 1 1 1 1 0 1 0 0 0 1 1 1 1 1 0 0 0 1 1 1 0 0 1 2 0 1

0 1 0 0 0 0 0 2 1 0 0 2

Melanorosaurus

1 0 0 1 ? 1 0 3 ? 0 1 2 1 1 1 1 1 1 0 1 ? 1 1 1 0 1 0 1 1 1 1 ? ? ?

0 1 0 1 0 1 1 0 1 0 1 1 0 0 1 1 0 1 0 1 1 ? 0 0 0 0 0 0 1 0 2 ? 0

1 1 0 1 1 0 0 1 ? ? 2 0 0 2 0 1 0 0 1 0 0 1 1 1 0 1 ? 0 0 1 0 1 ?

1 0 0 0 2 1 0 ? 1 0 0 2 1 1 0 0 1 1 0 ? 0 1 0 1 1 0 0 1 1 0 1 1 0

1 0 0 1 0 1 1 1 ? 0 0 0 0 0 0 1 ? ? 0 0 2 1 0 0 0 0 0 0 0 0 1 0 0

0 0 0 0 0 ? 0 2 0 1 0 0 1 1 0 0 1 0 ? 0 0 1 1 0 0 ? ? ? ? 0 0 0 ? ?

2 1 2 0 0 0 0 0 1 1 0 ? ? ? ? ? ? 0 0 ? 0 ? 2 1 0 0 1 0 1 1&2 0 ? 0

0 0 1 0 2 0 0 0 1 0 0 0 3 1 0 0 0 0 0 1 1 1 1 0 1 0 0 0 1 ? ? ? 1

? 0 1 0 1 1 0 ? 1 1 1 0 1 1 1 1 0 0 1 1 1 0 0 1 1 1 0 1 0 0 0 0 0

0 1 1 0 1 1 0 0 1 0 2 1 1 0 1 0 1 0 0 0 0 1 1 1 0 0 1 0 1 1 1 0

0 1 ? 1 1 1 1 1 0 0 0 0 1 1 0 ? 3

A. M. Yates28

Neosauropoda

1 1 0 0 0 1 1 3 1 1 1 2 0 1 1 0 1 2 0 0 0 0 0 1 0 1 0 1 1 1

0&2 1 1 1 1 1 1 2 0 1 0 1 0 1 0 1 1 0&1 1 1 1 1 0 1 2 0 1 0

0 0&1 1 0 1 1 2 2 2 1 1 1 0 0 1 0 1 1 1 2 0 0&1 2 0 0 0 0 0

1 0 2 0 1 1 1 0 0 0 0 1 2 0 1 0 0 0 0&2 1 1 1 1 1 1 0&2 1 2

0&1 1 1 2 0&1 0 0 0 0 1 1 1 0 0 1 1 1 1 0 1 0 0 1 1 0 0 2 0

1 2 1 0 1 0 1 0 1 0&1 0 2 0 1 1 1 0 0 1 1 0 2 0 1 2 1 1 0 0 0

0 2 1 0 0 1 1 1 0 0 1 1 1 1 0&1 1 0 0 0 0 1 0 0 1 1 1 0 0 3 1

1 0 0 1 0&1 0 0 1 1 1 1 0 0 0 1 0 1 0 0 0 0 0 0 0 0 1 1 0 0 0

1 ? ? 1 ? 3 0 1 1 1 1 0 0 3 1 0 2 1 0 0 0 0 1 1 1 1 0 0 1 1 0 0 1

1 1 0 0&1 1 0 0&1 0 0 2 1 1 1 1 2 ? ? 0 ? ? ? 0 1 0 ? 1 1 1 2 1

0 1 0 ? 1 1 0 0 0 0 1 1 1 1 1 0 1 0 1 1 1 1 1 1 0 1 1 ? ? 1 1 1 1

0 0 2 1 0 2 0 0 2 1 1 1 1 1 0 2 1 1 0 5

Neotheropoda

0 0 0&1 0 0&1 0 0 2 0&1 0 1 2 0 1 0 0 1 1 0 0 0 0 1

0&1 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 1 0 0&1 1

0 0 0&1 0 1 0 1 0 0 0 1 0&1 0 0 0 1 0&1 0 1 0 0 0 0 0

0 1 1 1 0 0 0 0 0 0 1 1 1 0 0 0 1 0 0 0 0 0 0 0 0 1 ? 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 1 1 0 0 0 0 0 1

0 0 0 0 0&1 1 0 0 0 0 1 0 2 0 0 0 0 1 0 1 0 1 1 0 0 1 0 0

0 0 0 0 1 0 0 0 0 0 0 0 0 0 1&2 1 0 0 0 1 1 0 0 0 0 0 0 0

0 0 1 0 1 0 0 0 0 1 0 0 1 0 1 1 0 0 0 0 0 1 0 0 0 1 0 0 0

1 2 0 0 0 0 0 0 1 0 ? 0 0 0 1 0 0 1 1 0 1 1&2 1 0 1 0 1 0

0 3 0 0 0 2 1 1 1 1 1 0 1 0 0 0 0 1 1 1 1 1 0 0 0 0 1 1 0

1 0 0 0 0 1 ? 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0&1 0 1 1 0 1

1 1 1 1 0 0 2 0 0 1 0 1 1 0 0 0 0 1 1 1 1 1 0 0 0 ? 0 ? 0 0

0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 1 2

Omeisaurus

1 1 0 0 0 1 1 3 1 1 1 2 0 1 1 0 1 2 0 0 0 0 0 1 0 1 0 1 1 ? 2

0 1 0 1 1 1 2 0 1 0 1 0 1 0 1 1 1 1 1 0 1 0 1 2 0 1 0 0 0 0 0

1 1 2 2 2 1 1 1 0 0 1 0 1 1 ? ? ? 0 ? 0 0 ? ? ? 1 ? ? ? 1 1 1 0 0

0 0 1 ? 0 ? 0 0 0 2 1 1 0 1 1 1 1 1 2 1 1 1 3 ? ? 0 0 1 1 1 0 1

2 0 0 1 1 0 1 0 1 1 1 0 0 2 ? 1 2 1 0 1 0 1 0 1 1 0 2 0 1 1 0

0 0 1 1 0 2 0 1 2 1 1 0 0 ? 0 2 1 ? ? 1 ? 1 0 0 1 1 1 0 0 1 0 0

? 0 1 1 1 1 1 1 0 0 3 1 1 0 0 1 0 0 0 1 1 ? ? ? ? ? ? 0 1 0 0 0

0 1 0 0 0 1 1 0 0 0 1 0 0 1 ? 3 0 1 1 1 1 0 0 3 1 0 1 1 0 0 0

0 1 1 1 1 0 0 1 1 0 0 0 1 1 1 1 0 0 1 0 0 2 1 1 0 1 2 ? ? 0 ? ?

? 0 1 0 ? 0 1 0 2 1 0 1 0 0 1 1 ? 0 ? ? 1 1 1 ? 0 1 1 0 1 0 ? 1

1 ? ? 1 ? ? ? ? 1 1 1 1 0 2 1 0 2 0 0 2 1 1 1 1 1 0 2 ? 1 0 5

Ornithischia

0 0 1 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 ? 0 0 0 ? 1 0 1 1 0 0

0 1 0 1 0 0 ? 1 0 0 0 0 0 0 1 0 0 1 0 0 0 0 1 0 1 0 0 0 0

0&1 0 0 0 1 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 1

0 0&1 0 1 0&1 0 0 0 0 0 0 1 0 0 1 1 1 0 0 1 1 1 0 1 0 0 0

0 1 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0&1 0 0 0 0 ? ? 0 1 0 0 0

0 0 0 0 ? 0 0 ? 0 ? 0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1&2 1

0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 1 0 1 1 1 1 1 0 0 0

0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 1 1 0 1 0 2 1 0 0 0 1 0 0 1 1 ? ? 0 0 0 ? 0 0 0 1 1 0 0

0 0 0 0 0 1 0 0 0 0 1 ? 0 1 0 1 1 1 1 0 2 0 0 0 0 0 0 0 0 0

0&1 0 1 1 1 1 0 0 0 2 ? 0 1 ? 0&1 1 0 0 0 0 0 1 1 1 1 0 0 0

0 0 0 0 0 0 0 0 0&1 1 0 0 0 0 0 0 0 0 0 0 1 0

Patagosaurus

1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? 1 ? 0 0 0 0 1 ? ? ? ? ? ? 0 ? ? 0 1 1 1 0 1 2

1 1 1 ? ? ? ? ? ? 1 1 0 0 1 0 1 1 0 0 0 0 0 1 1 0 0 2 0 ? 2 0 0

1 0 1 0 ? 0 0 2 0 1 ? 0 1 1 1 1 1 2 0 1 2 1 ? 0 0 ? 0 2 1 0 0 1

1 1 0 0 ? 1 ? 0 ? 1 0 0 ? ? ? 0 ? 1 1 1 0 0 ? 1 1 0 0 1 0 0 ? 1 1

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 1 1 0 0 3 1 0

1 0 0 0 0 0 1 1 1 1 0 0 1 1 0 0 1 1 1 1 1 0 0 1 0 ? 2 1 1 1 1

2 ? ? 0 ? ? ? 0 0 1 1 1 1 1 ? 1 0 1 0 0 0 ? ? 0 ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 5

Plateosaurus engelhardti

1 0 0 1 1 0 0 1 1 0 1 2 0 1 1 0&1 1 1 1 1 1 2 1 1 1 1 0 0 1

0 1 1 0 0 0 1 0 0 1 0 1 0 1 0 1 1 0 0 1 1 0 0 1 1 0 1 0 1 1 0

0 1 0 0 1 1 0 1 1 0 1 1 0 1 1 1 1 1 1 0 1 1 0 0 0 1 0 1 1 1 1

0 0 1 0 1 1 0 1 0 1 1 1 1 2 1 0 0 1 0 1 0 1 0 0 0 0 1 1 0 0 1

1 1 1 0 0 1 1 0 1 0 0 0&1 0 1 1 0 1 1 0 1 0 0 0 2 0 0 1 1 1

0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 1 0 0 0 1 0 0 0

0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 1 1 1 2 0 0 0 1 1 1 0 0 1 1

0 1 1 1 1 0 1 0 1 1 1 0 0 1 0 0 2 0 1 0 0 0 0 0 1 0 0 0 1 0 0

0 3 1 1 0 0 1 0 2 1 0 1 0 1 0 0 0 1 1 1 1 1 0 0 1 0 1 1 1 1 0

0 0 0 1 1 1 1 0 0 0 0 1 0 1 1 0 0 0 1 0 0 0 0 1 0 1 1 1 1 1 0

0 1 0 0 1 1 0 1 0 1 0 0 0 1 0 1 1 1 0 0 0 1 1 0 0 0 1 1 1 1 0

1 0 0 0 0 0 1 0 0 0 4

Plateosaurus gracilis

? 0 0 ? ? 0 0 1 ? 0 1 2 ? 1 1 0 1 1 1 ? 1 ? ? 1 1 1 0 ? 1 0 1 ? ?

0 ? 1 0 ? 1 0 1 0 ? 0 ? ? 0 ? 1 0 0 0 1 1 0 1 ? ? 1 0 ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? 0 1 1 0 ? ? ? ? ? 1

2 1 0 0 1 0 1 0 1 0 ? 0 0 1 ? ? ? ? ? 1 1 0 0 ? ? 0 1 ? ? ? 0 1 1

0 1 1 0 1 ? 0 0 ? 0 0 1 ? 1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0

0 1 0 ? 1 0 1 0 0 0 1 0 1 0 0 0 0 0 0 1 0 0 0 ? 0 0 ? ? ? ? ? ? ?

1 2 0 0 ? 1 1 1 0 0 ? 1 0 1 1 ? 1 0 1 0 1 0 1 0 0 0 0 ? ? ? 1 0

? ? ? ? ? ? 0 0 1 0 0 0 3 1 1 0 0 1 0 2 1 0 1 0 1 0 0 0 1 1 1 1

1 0 0 1 0 1 1 ? ? 0 0 0 0 1 1 1 1 0 0 0 0 1 0 1 1 0 ? 0 1 0 0

0 0 ? 0 ? ? 0 1 1 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? ? ? ? ?

? ? ? 0 ? 0 ? ? ? ? ? ? ? ? 2&3

Plateosauravus

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? 1 0 ? ? ? 0 1 0 0 1 0 1 ? 0 1 1 0 1 ? 0 0 1/2 0 0 1 1 ? 0

0 1 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 ? ? 1 0 ? ? 0 1 1 ? ? ? 0

? ? 1 1 0 ? ? ? ? ? ? ? ? 0 ? ? 2/3 1 1 1 0 0 1 1 1 0 0 ? ? ? ? ? ? ?


? ? ? ? ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 1 0 0 0 3 1 1 0 0 1 0 1

? 0 1 ? ? ? ? ? ? ? ? 1 1 ? ? 1 0 1 1 0 ? 0 0 0 0 1 1 1 1 0 0 0 0

1 1 1 1 0 0 0 1 0 0 0 0 0 0 1 1 0 1 1 0 ? ? ? ? ? ? 0 ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? 1 1 ? 0 ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? 4

Riojasaurus

1 0 0 1 ? 0 0 ? ? 0 1 2 0 1 1 ? 1 1 0 ? ? 0 0 1 1 1 0 1 1 0

1 1 0 0 ? 1 0 0 0 0 1 0 0 0 ? 1 0 0 1 0 ? 0 0 1 1 1 0 0 1 0

0 0 0 0 1 1 0 1 ? 0 ? 1 0 0 1 1 ? 0 1 0 1 1 ? ? ? ? ? ? 1 ? 0

0 1 0 0 0 ? 1 ? ? 1 0 1 ? 2 0 0 0 1 0 1 0 1 0 0 0 0 1 0 ? ?

1 1 1 1 0 0 1 1 0 1 0 0 1 0 1 1 0 1 1 0 1 0 0 0 1 0 0 1 1

1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 1 0 1 0 0 1

0 0 0 0 0 0 0 1 1 0 0 0 ? ? 0 ? 0 1 0 1 1 2 1 2 1 0 1 1 1 1

0 0 1 1 1 1 1 1 ? 0 1 1 1 1 1 0 1 1 0 0 1 0 1 0 0 0 1 0 2 ?

0 0 1 0 0 0 3 1 1 0 0 1 0 2 1 1 1 0 1 0 0 0 1 1 0 1 1 0 0

1 0 1 1 0 0 1 0 1 1 0 1 1 1 0 1 0 1 1 0 2 0 1 1 0 1 0 0 0

0 0 0 1 1 0 1 1 0 0 1 0 0 1 1 0 1 0 1 0 0 ? ? 0 1 1 ? 0 0 0

1 1 1 0 0 1 2 1 1 1 1 0 0 0 0 0 1 ? ? 1 3

Ruehleia

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? 1 0 ? ? ? 0 ? ? ? ? 0 1 ? ? ? 1 0 1 ? 0 0 2 0 0 1 ? ? 0 0 1

1 0 0 1 0 0 0 0 0 1 1 0 0 0 0 0 0 ? 1 1 0 0 0 0 1 ? 0 0 ? 0 ? 0

1 1 0 ? ? ? ? ? ? 0 1 0 1 1 1 1 2 0 0 0 1 1 1 0 0 ? 0 1 0 0 ? ? 0

1 0 1 1 1 0 1 0 0 0 ? ? ? 0 ? ? 0 0 ? ? 0 0 1 0 0 0 3 1 1 0 0 0

0 2 1 ? ? 1 1 0 0 0 1 1 0 1 1 0 ? 1 0 1 1 0 ? 1 0 1 0 1 1 0 1 0

0 0 0 1 0 2 1 0 0 0 1 0 ? 0 0 ? 0 1 1 0 1 1 0 0 1 ? 0 0 0 0 1

0 1 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 3

Saturnalia

1 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 ? ? 1 ? ? ? ? 0 ? 1 0

0 ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 0 ? ? ? ? ? 1 ? ? 0 0 ? ? ? 0

? ? ? 0 ? 0 0 0 0 ? ? ? ? ? ? ? ? ? ? 0 1 0 0 0 ? ? ? ? ? ? 0 0 0 0 1

0 0 0 1 0 0 0 0 ? ? ? ? ? ? ? ? 0 0 ? ? 0 1 0 0 1 0 ? ? 0 1 1 0 1 ?

0 0 0 0 0 1 ? 0 0 0 1 1 0 0 1 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0

0 0 0 0 1 0 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 1 1 1 1 1 1 0 0 0 1 0

1 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? 0 ? 1 1

1 0 0 2 1 0 2 1 0 0 0 0 0 0 0 1 0 0 1 1 0 0 1 0 0 1 0 0 0 0 0

0 1 0 0 1 1 0 0 0 1 0 1 1 0 0 0 1 0 0 0 0 0 0 0 1 0 1 0 0 0 1

0 0 1 0 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0

? 0 0 0 ? 0 0 1 0

Shunosaurus

1 1 0 0 0 1 1 3 ? 1 1 2 0 1 1 0 1 2 0 0 0 0 0 0 0 1 0 1 1 ? 2

0 1 0 1 1 1 2 0 1 0 1 0 1 0 1 0 0 1 1 0 1 0 1 2 1 0 0 0 1 1 0

1 1 1 2 1 0 1 0 0 0 1 0 1 ? 1 2 0 0 2 0 ? ? ? 1 1 0 2 0 0 1 0 0

0 0 0 1 2 0 1 0 0 0 1 1 1 1 1 1 1 2 1 2 1 1 1 2 ? ? 0 1 0 1 1

? 1 0 0 1 ? 1 ? 1 0 0 1 0 0 1 2 ? 1 1 0 0 0 0 1 0 1 1 0 2 1 ? 0

? 0 0 1 ? 0 2 0 1 2 1 0 0 0 0 ? 1 1 ? 0 0 ? 1 0 0 1 1 1 0 1 1 0

0 1 ? 1 1 1 ? 1 ? 0 0 2 1 1 0 0 1 1 0 0 ? 1 1 0 0 0 ? 1 0 1 1 0

0 1 ? 0 0 0 1 1 0 0 0 0 ? ? 1 ? 3 0 1 1 1 1 0 0 3 1 0 1 1 0 0 0

0 1 1 1 1 ? 0 1 1 0 0 ? 1 1 1 1 0 0 1 0 0 2 1 1 1 1 2 ? ? 0 ? ?

? 0 0 0 ? 0 1 1 2 1 0 1 0 ? 0 1 ? 0 ? ? ? ? 1 1 0 ? 1 0 1 ? ? ? 1

1 0 1 1 ? ? 1 1 1 1 1 0 2 0 0 2 ? 0 2 1 1 1 ? 1 0 2 1 1 0 5

Silesaurus

0 0 ? 0 ? ? 0 0 ? 0 0 0 ? 0 0 1 0 0 0 0 0 ? ? 0 ? 0 0 ? 0 ? 0 1 0

1 ? ? ? ? ? ? ? ? ? 0 0 1 0 ? ? 0 ? ? ? ? 0 1 ? ? ? ? ? ? ? ? ? 0 0 ? 0

? 0 ? 0 0 0 ? ? 0 0 0 0 0 0 ? ? ? ? ? 1 ? 0 0 ? 0 ? 0 1 0 ? ? ? 0 ?

0 0 0 0 0 1 0 0 0 1 0 0 0 0 1 0 ? 0 0 0 0 0 0 0 0 0 0 0 ? ? ? 0

0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0

0 0 0 0 0 ? ? 0 0 0 0 0 1 0 0 0 0 0 ? 0 ? 0 ? ? ? ? ? ? ? 1 1 1 1

1 1 0 0 0 0 1 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 ? 0 1 0

1 0 0 ? 0 ? 1 0 0 0 0 1 1 0 1 1 0 0 0 1 0 1 1 0 0 0 1 0 0 0 1

1 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 ? ? 1 1 ? 1 ? ? 0 0 ? ? 0 0

0 0 0 0 ? 1 0 ? ? ? 0 0 0 ? 0 0 1 1

Staurikosaurus

0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? 0 0 0 0 0 0 0 0 0 1 0 0 0 ? 0 0 ? 0 0 0 0 0 0

0 0 0 0 ? ? ? ? ? ? ? ? 0 0 0 1 0 ? ? ? ? 0 0 ? 0 1 0 0 1 ? 0 0 0 0

1 1 ? 0 0 0 ? 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 1 ? ? 0 1 0 1 0 1

1 0 0 0 0 0 1 1 0 0 1 0 ? ? ? ? 1 ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 0 0 0 1 2 0 0 1 1 0

0 0 1 1 1 0 ? 0 1 0 ? 0 0 0 1 0 ? 0 0 0 0 0 1 0 0 0 0 1 ? ? 0 0

0 ? 0 1 0 1 ? 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 ? ? ? ? 0 1 ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1

Tazoudasaurus

1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? 0 1 0 ? ? ? ? ? ? ? ? ? ? ? ? 0 0 ? 1 ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 0 0 0 0 0 1 ? ? ? 0 ? ? 2 0 ? 1 1 1 1 0

1 2 1 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 2 ? ? 1 ? ? 0 0 1

? ? 1 0 2 0 1 1 ? 0 0 1 ? ? 2 0 1 1 1 ? 0 0 ? ? ? ? ? ? ? ? 0 0 ? ?

? ? 1 1 0 ? ? ? ? 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 1 0 1 0

0 0 1 ? ? ? ? ? ? ? ? ? ? ? ? 1&2 1 ? ? ? 1 ? 1 0 0 1 ? 0 ? ? ? ? ? ?

? 0 ? ? ? ? 0 ? ? ? ? ? ? ? ? ? 1 0 1 0 1 ? 1 1 1 ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? 1 ? ? 1 0 ? 1 ? ? ? ? 5

Thecodontosaurus antiquus

? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 ? ? 1 ? ? 1 0 0 ? ? ? ?

? ? 0 ? 0 ? 0 ? 0 ? ? ? 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? 1 0 ? 0 ? 0 ? 1 0

1 ? ? 0 1 1 0 0 0 ? ? ? ? ? ? ? 0 0 ? 1 1 0 1 0 ? ? ? ? ? ? 2 1 0 0

1 0 1 0 1 0 0 0 0 ? 1 ? ? ? ? ? ? 0 0 0 1 0 1 0 1 0 ? 1 ? 0 1 1 0

1 ? 0 0 0 0 0 1 ? ? 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 1

A. M. Yates30

0 0 0 ? 0 1 0 1 1 0 0 ? 0 1 0 0 0 ? ? 0 0 ? 0 1 0 1 1 2 1 1 0 0

0 1 0 1 0 0 1 1 0 0 0 1 2 0 1 0 0 0 1 1 0 0 0 0 1 0 1 0 0 0 0

0 1 1 0 0 0 0 0 0 2 1 0 0 0 0 0 1 ? ? ? 0 ? 0 0 0 ? ? 0 1 1 ? ?

0 1 0 1 0 ? 0 0 0 0 1 1 1 1 0 0 0 0 1 0 1 1 0 0 0 1 0 0 0 0 0

0 0 1 0 1 1 0 0 ? ? 0 1 1 0 1 0 1 0 0 ? ? ? ? 1 1 0 0 0 1 1 1 0

? 1 ? 0 1 0 ? ? 0 0 ? 0 ? ? ? ? 1

Thecodontosaurus caducus

? 0 ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? 0&1 0 0 ? 0 ? ? ? 1 ? ? 1 ? ? ? ? 0

0 1 0 0 1 0 0 0 0 0 0 1 0 ? 1 0 0 0 0 1 ? 1 0 ? ? ? ? ? ? ? ? 0 0

? 0 0 0 ? 1 ? 1 ? ? 0 0 1 0 0 0 1 1 ? 0 0 0 ? 0 0 ? 0 1 0 1 0 ? ?

0 0 0 0 ? 1 0 0 1 0 1 0 1 0 0 0 0 1 0 0 1 ? ? 1 1 0 0 ? 1 0 1 0

1 0 0 1 1 0 1 1 0 1 ? 0 0 ? ? ? 1 ? ? ? ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? 0 ? ? 1 1 ? 0 0 0 1 0 0 0 0 1 0 0 ? ? ? ? 1 1 ? 1 ?

0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0

1 0 0 0 2 1 1 0 0 0 0 1 ? ? ? ? ? ? ? ? ? 0 0 1 1 ? ? 1 0 1 1 0 ?

0 0 ? ? ? ? ? ? ? ? ? ? 1 ? 1 ? 0 0 0 ? 0 0 0 0 0 0 0 ? 0 1 ? 0 0 1

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 1 1 1 0 0 1 1 ? 1 0 0 0 0 0 0 0

1 0 0 1 ?

Unaysaurus

1 0 0 ? 1 0 0 1 ? ? 1 ? 0 1 ? 1 1 1 ? ? ? 2 1 1 1 1 0 ? 1 0 1 1 0

0 ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? 0 0 0 0 1 ? 1 0 1 1 0 0 0 0 0 ? ? 0 ?

1 0 ? ? 0 ? 1 ? ? 0 1 0 1 0 0 ? ? ? ? ? ? ? 1 0 ? 1 0 1 1 0 ? ? ? ? ?

0 2 ? 0 0 1 0 1 0 1 0 0 0 0 ? ? ? 0 ? 1 ? ? ? ? ? 0 ? 1 ? 0 ? ? ? ?

? ? ? ? ? ? 0 ? ? 0 ? 1 ? ? 0 0 ? 1 0 0 0 0 ? 0 ? ? 0 0 0 0 ? ? ? ? ?

? ? ? ? ? ? ? ? ? 0 ? ? ? ? 0 1 0 0 0 ? 0 0 ? 0 1 0 1 1 ? 1 2 0 0 0

0 1 1 0 0 ? ? ? ? ? ? 1 ? ? ? ? 1 1 0 ? ? ? ? 2 1 1 ? ? ? 0 ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 0 1 1 0 ? ? ? 0 1 1 0 1 0 ? ? ?

? ? ? ? ? ? ? 0 ? 1 1 0 ? ? ? 0 ? ? 0 ? ? ? ? ? ? ? ? ? ? 1

Vulcanodon

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1/2 ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 ? ? ? ? ? ? 0 1 1 1 ? 0 ? 0 0 ? 0 ? 0 ?

? 0 1 ? ? 2 1 1 0 ? 1 0 ? 0 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

? ? ? ? ? ? ? ? ? ? ? ? ? ? 3 1 0 1&2 ? 0 0 ? 1 1 1 1 1 0 0 0 1 0

0 1 1 1 1 1 0 0 1 0 ? 1 1 1 0 1 1 1 1 0 0 1 0 0 1 1 ? 1 ? ? 2 0

? 1 0 0 0 1 ? 0 ? ? ? 1 1 ? 1 0 1 0 1 1 0 1 1 1 0 1 1 ? ? 1 1 ? 0

1 0 1 0 1 2 0 0 1 1 1 1 0 ? 1 2 1 ? 0 5

Yunnanosaurus

1 1 0 ? ? 0 0 2 ? 0 1 ? 1 1 1 1 1 1 0 ? ? ? ? 1 0 1 ? 1 1 ? 0 1 0

1 ? 1 0 1&2 ? ? ? ? ? 0 ? ? 0 ? ? ? 0 1 1 1 ? 0 0 ? ? 0 0 0 1 0 ?

? 0 1 ? 0 ? ? 0 0 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 0 ? 0 0 0 1 0 ? ? ?

0 ? 0 1 1 0 0 1 0 1 2 1 2 1 1 1 1 ? ? ? 1 1 1 1 0 0 1 1 0 1 1 0

1 0 0 ? 0 1 1 0 1 0 0 0 0 0 0 1 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0

1 0 0 0 0 0 0 ? 0 1 0 1 1 0 1 0 0 0 0 0 ? 0 0 ? 0 ? ? 0 ? ? 0 0

1 ? ? ? 1 1 2 0 0 ? 1 1 1 0 0 ? ? 0 1 ? ? 0 0 1 1 ? 2 1 0 1 1 0 0

2 0 1 0 0 0 0&1 0 2 ? 0 0 1 0 0 0 3 1 0 0 0 0 0 0 1 1 1 0 1

0 0 0 1 0 ? 1 1 0 0 1 0 1 1 1 ? 0 0 1 0 1 1 0 1 0 0 0 0 1 0 1

1 0 1 0 1 0 1 0 0 0 0 1 0 0 1 1 0 0 1 ? 0 1 1 0 1 0 1 0 0 0 1

1 1 1 0 0 1 0 1 1 1 0 0 1 ? ? ? 1 1 ? 0 0 0 0 1 1 0 ? 2


Yates

Documents

Transcript of Yates