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Ankylosaurs from the Price River Quarries, Cedar Mountain Formation (Lower Cretaceous), East-Central Utah

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A new large nodosaurid ankylosaur, Peloroplites cedrimontanus, is described from a partial skull and postcranial skeleton found at the PR-2 Quarry located at the base of the Mussentuchit Member of the Cedar Mountain Formation in central Utah. The specimen is about the same size as the contemporary nodosaurid Sauropelta edwardsorum from the Cloverly Formation of Montana, and is of an individual approximately 5–5.5 m long. The skull of Peloroplites differs from that of Sauropelta in the vertical orientation of the suspensorium, non-domed cranium and broad, square premaxillary beak. The quarry is near and roughly at the same level as the CEM Quarry that produced the holotype of the ankylosaurid Cedarpelta bilbeyhallorum. The postcrania of Cedarpelta is described and illustrated based on the paratype and new material. These elements clearly establish that Cedarpelta is closer to Ankylosaurus than to Sauropelta. As a primitive ankylosaurid, there is no a priori reason to assume that the tail club was present. Based on recent finds in China, a hypothesis is presented that the tail club is a derived feature in non-shamosaurine (i.e., ankylosaurine) ankylosaurids.
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ARTICLE
ANKYLOSAURS FROM THE PRICE RIVER QUARRIES, CEDAR MOUNTAIN FORMATION
(LOWER CRETACEOUS), EAST-CENTRAL UTAH
KENNETH CARPENTER,
*,1
JEFF BARTLETT,
2
JOHN BIRD,
2
and REESE BARRICK
2
1
Department of Earth Sciences, Denver Museum of Nature & Science, 2001 Colorado Blvd., Denver, CO 80205;
Ken.Carpenter@dmns.org
2
Prehistoric Museum, College of Eastern Utah, Price, UT 84501
ABSTRACT—A new large nodosaurid ankylosaur, Peloroplites cedrimontanus, is described from a partial skull and
postcranial skeleton found at the PR-2 Quarry located at the base of the Mussentuchit Member of the Cedar Mountain
Formation in central Utah. The specimen is about the same size as the contemporary nodosaurid Sauropelta edwardsorum
from the Cloverly Formation of Montana, and is of an individual approximately 5–5.5 m long. The skull of Peloroplites
differs from that of Sauropelta in the vertical orientation of the suspensorium, non-domed cranium and broad, square
premaxillary beak. The quarry is near and roughly at the same level as the CEM Quarry that produced the holotype of
the ankylosaurid Cedarpelta bilbeyhallorum. The postcrania of Cedarpelta is described and illustrated based on the
paratype and new material. These elements clearly establish that Cedarpelta is closer to Ankylosaurus than to Sauropelta.
As a primitive ankylosaurid, there is no a priori reason to assume that the tail club was present. Based on recent finds in
China, a hypothesis is presented that the tail club is a derived feature in non-shamosaurine (i.e., ankylosaurine) anky-
losaurids.
INTRODUCTION
The past decade has seen the discovery of numerous new an-
kylosaur taxa globally. These discoveries have shed much new
light on ankylosaur evolution and relationships, while at the
same time creating many new questions. The once clear di-
chotomy of two families, the Nodosauridae and Ankylosauridae
(Coombs, 1978), is now blurred with the recognition of a third
family, the Polacanthidae (Carpenter, 2001) based on ankylo-
saurs from the Jurassic (Carpenter et al., 1998) and Lower Cre-
taceous (Kirkland, 1998). Furthermore, the Ankylosauridae is
now subdivided into the subfamilies Shamosaurinae and Anky-
losaurinae (Tumanova, 1983; Vickaryous et al., 2001). The Po-
lacanthidae is not yet universally accepted (e.g., Vickaryous et
al., 2004), but it has been independently substantiated by its
distinctive armor histology (Scheyer and Sander 2004).
Ankylosaurs from the Lower Cretaceous Cedar Mountain
Formation of eastern Utah (Fig. 1) have proven to be uncom-
monly rich and diverse compared to similar aged strata. Whereas
most Lower and even middle Cretaceous strata from North
America only contain a single ankylosaur taxon each (Carpenter
and Kirkland, 1998), the Cedar Mountain Formation has at least
six, probably more (Carpenter et al., 1999, 2002). These ankylo-
saurs are not evenly distributed throughout the formation, how-
ever. Polacanthid ankylosaurs (Gastonia burgei, Gastonia n.sp.
and large unnamed taxon, Kirkland, 1998; Carpenter et al., 2007;
Kirkland and Madsen, 2007) occur in the Yellow Cat through to
basal Ruby Ranch Members, thus apparently become extinct
during the earliest Aptian. The oldest ankylosaurid, Cedarpelta
bilbeyhallorum appears at the base of the Mussentuchit Member
and was briefly described by Carpenter et al. (2001). The oldest
nodosaurid is an unnamed large taxon that appears about in the
middle of the Ruby Ranch (Warren and Carpenter, 2004) and
the oldest named taxon is Animantarx ramaljonesi from low in
the Mussentuchit Member. A previous report of cf. Sauropelta
sp. from the older Poison Strip Member by Carpenter et al.
(1999), following Kirkland (1992, 1996), is probably in error. The
specimen may in fact belong to a new, large polacanthid as dis-
cussed further below.
A large nodosaurid from the Cedar Mountain Formation was
previously mentioned by Burge and Bird (2001). More of that
nodosaurid is now known and is named below. The new taxon
comes from the Price River II quarry (a.k.a., PR-2), which is
about 24.5 km southeast of Price, Emery County, Utah (Fig. 1).
PR-2 has been reported previously as occurring in the Ruby
Ranch Member (Burge and Bird, 2001; Burge et al., 2000), but
the dark, carbonaceous nature of mudstones identifies the strata
as the Mussentuchit Member. In addition to the new nodosaurid,
the quarry has produced four individuals of a new brachiosaurid
sauropod, an iguanodontid, material referable to the ankylosau-
rid Cedarpelta bilbeyhallorum (described below), a turtle (Burge
and Bird, 2001; Burge et al., 2000; Coulson et al., 2004), and a
pterosaur. Radiometric dates (109–116 Ma) place the quarry at
the Aptian-Albian boundary (Burton et al., 2006) based on the
2004 revised geologic time scale (Gradstein and Ogg, 2004).
Thus, this part of the Cedar Mountain Formation is equivalent
with at least part of the Cloverly Formation of Wyoming and
Montana (108.5 +/− 0.2 Ma, Burton et al., 2006).
Institutional AbbreviationsAMNH, American Museum of
Natural History, New York, New York; CEUM, College of East-
ern Utah, Prehistoric Museum, Price, UT; GISPS, Geological
Institute, Section of Paleontology and Stratigraphy, Ulan Bataar,
Mongolia; HMNS, Hayashibara Museum of Natural Sciences,
Okayama, Japan; KUVP, Kansas University, Vertebrate Pale-
ontology, Lawrence, Kansas; SMU, Southern Methodist Univer-
sity, Dallas, Texas; USNM, United States National Museum
(now the National Museum of Natural History), Washington,
D.C.
*
Corresponding author.
Journal of Vertebrate Paleontology 28(4):1089–1101, December 2008
© 2008 by the Society of Vertebrate Paleontology
1089
Comparative MaterialsComparisons are made with the po-
lacanthid Gastonia (CEUM 1307, and thousands of bones at the
College of Eastern Utah Prehistoric Museum and Denver Mu-
seum of Nature & Science (ms in preparation); nodosaurids Ani-
mantarx (CEUM 6228), Edmontonia (AMNH 5581; HMNS-001;
USNM 11868), Pawpawsaurus (SMU 73203), Sauropelta
(AMNH 3032), and Silvisaurus (KUVP 10296); ankylosaurids
Ankylosaurus (AMNH 5214), Euoplocephalus (AMNH 5404),
and Saichania (GISPS100/1305).
SYSTEMATIC PALEONTOLOGY
Order ANKYLOSAURIA Osborn 1923
Family NODOSAURIDAE Marsh 1890
PELOROPLITES, gen. nov.
Etymology—“monstrous heavy onederived from the Greek
peloros monstrous, gigantic; and hoplites, meaning heavily
armed; and as a subjunctive, a heavily armed soldier.
Type SpeciesPeloroplites cedrimontanus.
Diagnosisas for the type and only species.
PELOROPLITES CEDRIMONTANUS, sp. nov.
EtymologyCedrus Latin for cedarand mont- Latin com-
bining form of mountain, meaning from Cedar Mountainin
reference to the formation.
Type Locality and StratumPrice River II Quarry (Locality
number EM 372), base of the Mussentuchit Member, Cedar
Mountain Formation, Emery County, Utah. Exact locality infor-
mation is available from the Prehistoric Museum, College of
Eastern Utah.
DiagnosisSkull lacking premaxillary teeth, occiput slopes
anterodorsally, no prominent lateral temporal notch dorsally as
in Sauropelta; cranial hornssmall, and blunt, paroccipital pro-
cess project laterally, not posterolaterally as in Sauropelta; quad-
rate vertical, not anteriorly bowed or sloped anteroventrally;
odontoid very short, unlike Sauropelta, axis centrum short, as
long as tall. Coracoid:scapula proportions about 2:3, similar to
Animantarx,Edmontonia, unlike 1:3 in Sauropelta.
HolotypePartial skull CEUM 26331.
ParatypesCervicals CEUM11342, CEUM 11354, CEUM
11682, CEUM 26283, CEUM 36714; dorsals: CEUM 8080,
CEUM 11355, CEUM 11654, CEUM 11702, CEUM 11720,
CEU12829, CEUM 36701; synsacrum CEUM 50418, CEUM
11889; caudals: CEUM 19481, CEUM 22684, CEUM 31961,
CEUM 31971, CEUM 32003, CEUM 32022, CEUM 36696,
CEUM 11330, CEUM 36703, CEUM 36709, CEUM 39582,
CEUM 50410, CEUM 11624, CEUM 11681, CEUM 11632,
CEUM 12755, CEUM 12846, CEUM 14382, CEUM 36186;
Chevron CEUM 11735; fragments; CEUM 19507, CEUM 11670,
CEUM 14381. Scapula-coracoid: CEU11706, CEUM 11719; hu-
merus: right CEUM 10614, CEUM 11334, right CEUM 11639,
CEUM 12831; radii: CEUM 11631, CEUM 11655, CEUM 11712,
CEUM 11718; ulnae: CEUM 11708, CEUM 11347, CEUM
11888, CEUM 11669; ilia: CEUM 26342, right CEUM 50427;
pubis CEUM 11626; ischium: distal end CEUM 11313; Femora:
head CEUM 22682, CEUM 11319, CEUM 11321, CEUM 11676;
Tibiae: fragment CEUM 12837, CEUM 11640, CEUM 11710;
fibula: CEUM 12836, CEUM 12158; metacarpals: CEUM 12187,
CEUM 12188, CEUM 12189, CEUM 12191, CEUM 12193;
metatarsal: CEUM 11605; metapodials: CEUM 10638, CEUM
11606, fragment CEUM 11617, CEUM 11639, CEUM 11641,
CEUM 11649, fragment CEUM 11665, CEUM 11666, CEUM
11672, CEUM 11673, CEUM 11700, CEUM 14390; fragment
CEUM 26286 CEUM 19527, CEUM 31239, CEUM 35866; pha-
langes: CEUM 11357, CEUM 11618, CEUM 11623, CEUM
11882, CEUM 11646, CEUM 11656, CEUM 12190, CEUM
12192, CEUM 12219, CEUM 12220, CEUM 12222, CEUM
12834, CEUM 20562, CEUM 32011, CEUM 50383; unguals:
CEU11645, CEUM 11647, CEUM 12164, CEUM 11671, CEUM
12218, CEUM 12221, CEUM 12223, CEUM 12797, CEUM
13286, CEUM 26324, CEUM 19553, CEUM 32002; armor:
CEUM 1344, CEUM 11971, CEUM 11346, CEUM 11350,
CEUM 11973, CEUM 11356, CEUM 11607, CEUM 11650,
CEUM 11697, CEUM 12825, CEUM 36404, CEUM 14050,
CEUM 12388, CEUM 12867, CEUM 26291, CEUM 20561,
CEUM 26333, CEUM 34578, CEUM 38748, CEUM 14331.
DESCRIPTION
Cranial
A reconstructed skull is shown in Figure 2. It is estimated to
have been 56 cm long and had a maximum width between the
dorsal orbital rims of 35.5 cm, which is about the same width as
Sauropelta AMNH 3035 (Carpenter and Kirkland, 1998). The
FIGURE 1. Stratigraphic section (A) and location map (B)ofPR2
relative to the Cedar Mountain outcrop belt (black) in eastern Utah.
Modified from Kirkland and Madsen (2007). Naturita Formation is used
in place of Dakota Formation for the reasons given by Witzke and Lud-
vigson (1994: 69-73). Naturita Formation, introduced by Young (1960,
1965), is the next available name for these strata.
JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 28, NO. 4, 20081090
snout of Peloroplites tapers anteriorly and terminates abruptly at
a relatively broad premaxillary beak, as compared to Silvisaurus.
The premaxillae are fused along their mid-line (Fig. 3AC).
Although the left premaxilla is damaged along its lateral margin,
the width of the premaxillary beak is estimated to be 18 cm. The
premaxillae are dorsoventrally thick, whereas they are thin in
Gastonia (unknown in Animantarx). The dorsal surface of the
premaxillae is rugose for the keratinous beak (smooth in Gasto-
nia), as well as arched in anterior view; in addition, the beak has
a broad, inverted U-shaped notch that is less developed than in
Gastonia. A groove is present near the lower margins of the beak
in anterior view (Fig. 3C). This groove continues on to the pala-
tal side (Fig. 3B) defining the lateral edge of the tomial ridge.
This ridge is inset from the lateral margins and the beak is con-
cave between them.
Both the left and right prefrontals and lachrymals are fused
(Fig. 4). The presence of the lachrymal is inferred from the lach-
rymal foramen seen within the anterior orbital wall (Fig. 4F, L).
Both sets of prefrontal-lachrymals are triangular in lateral and
dorsal views. Their external surfaces have rugose sculpturing
composed of irregular pits, anastomosing valleysand ridges.
This sculpturing is especially prominent over the orbits. A faint,
shallow groove is present just anterior to the orbit, which extends
onto the dorsal surface of the prefrontal. Similar grooves are
seen in other nodosaurids, such as Edmontonia, and probably
delineate the margins between adjacent keratinous scales in the
living animal. Medially, the prefrontal-lachrymals are divided by
the anterior orbital wall, which separates the orbit from the nasal
cavity. The wall is pierced by the lachrymal fenestra. The dorsal
surface of the orbit is concave.
The postorbital, squamosal, jugal, quadratojugal, and quadrate
are coossified on both side of the skull and all traces of sutures
are missing (Figs. 5, 6). The identity of the elements is inferred
based on topography relative to the landmarks of the orbit and
lateral temporal fenestra. In addition, the right complex is at-
tached to the braincase and dorsal part of the skull. The external
surfaces are rough and pitted, although not to the extent seen
over the orbits. The postorbital horncores (dorsal coronux of
Blows, 2001) are very low, conical structures that project dorso-
laterally. They are much less prominent than seen in many no-
dosaurids, such as Pawpawsaurus or Sauropelta, or even in the
polacanthid Gastonia The jugal-quadratojugal horncores (ven-
tral coronux of Blows, 2001) are not prominent, appearing as
low, localized thickening of bone; they are very prominent and
tapering in the polacanthid Gastonia, as well as in other nodo-
saurids, such as Pawpawsaurus and Animantarx, but moderately
developed in Sauropelta,Edmontonia and Silvisaurus.
The ventral rim of the orbit is probably composed of the jugal
(sutures have been obliterated). It forms a medial-lateral narrow
floor to the orbit. The posterior rim of the orbit is assumed to be
FIGURE 2. Reconstructed skull of Peloroplites cedrimontanus CEUM
26331. A, lateral; B, dorsal; C, posterior. Scale in cm.
FIGURE 3. Premaxillary beak of Peloroplites cedrimontanus CEUM
26331.1: A, dorsal; B, palatal; C, anterior views. Maxillary fragment with
tooth CEUM 26331.6: D, medial side showing tooth; E, lateral side; F,
close-up of tooth. Abbreviations:g, grove; n, notch; tr, triturating ridge.
Scale in cm.
CARPENTER ET AL.CEDAR MOUNTAIN ANKYLOSAURS 1091
composed of the postorbital and jugal as it typically does in other
dinosaurs. The posterior orbital wall is more complete on the
right side and shows that the wall did not completely separate the
orbit from the temporal fossa. Laterally on the skull, the orbit is
widely separated from the lateral temporal fenestra (Fig. 5A) as
in Edmontonia and unlike Pawpawsaurus. This great width sug-
gests a broad postorbital and jugal, but this needs to be con-
firmed from a younger specimen where sutures are visible. The
squamosal is fused to the head of the quadrate, although crush-
ing of the left side has displaced the head medially some. The
quadrates are slightly bowed anteriorly (seen best in Fig. 6A),
whereas they are more strongly bowed in Edmontonia,Animan-
tarx and Pawpawsaurus, and moderately bowed in Silvisaurus.
They are not inset from the lateral sides of the skull, as they are
in most nodosaurids, including Sauropelta (compare Fig. 6F with
Carpenter and Kirkland, 1998, fig. 9A). In these other nodosau-
rids, the inset of the quadrate recesses the lateral temporal fe-
nestra from the lateral margin of the skull and is manifest in
dorsal view as the lateral temporal notch (Carpenter and
Kirkland 1998). The lateral temporal fenestra, best seen on the
right side, is a vertical rectangle with rounded corners and is
almost keyhole shape.
The frontoparietal region is slightly domed in contrast to the
more extreme condition seen in Panoplosaurus (Carpenter,
1990) and possibly Sauropelta. In posterior or occiput view (Fig.
6C), the dome is moderately arched laterally in a manner similar
to Animantarx,Edmontona and Silvisaurus. The nuchal margin
is scalloped and overhangs the sutural region to the paroccipital
process and supraoccipital. The paroccipital process is twisted
along its length so that the posterior surface faces obliquely
downward. In Edmontonia and Animantarx the entire paroccipi-
tal faces obliquely downward and in Pawpawsaurus it faces
mostly posteriorly. The supraoccipital crest is weakly developed
as is typical for nodosaurids. The facet for the proatlas is seen on
the right side, although it is partially damaged. The exoccipital is
damaged on the left side and the exoccipital-basioccipital suture
of the right side is fused. The amount the exoccipital contributes
to the occipital condyle cannot thus be determined. The foramen
magnum is crushed somewhat so that its exact shape cannot be
determined. The occipital condyle has the typical hemispherical
shape of nodosaurids. In addition, the dorsal surface of the con-
dyle neck is slightly concave as in Pawpawsaurus.
Ventrally, the basioccipital-basisphenoid suture is fused and
assuming that the basitubera marks the approximate location of
the suture, the basioccipital is twice as long as the basisphenoid.
The lateral wall of the braincase is damaged making identifica-
tion of the fenestra somewhat problematic, and therefore we
refrain from doing so at this time. The parasphenoid extends
anterodorsally from the basisphenoid (Fig. 6B) and is fused to
the underside of the cranium. A fragment of the displaced pos-
terior pterygoid plate is present anterior to the parasphenoid
(Fig. 6D, E). The fragment is concave as in Edmontonia and
probably represents the left pterygoid. The adductor fossa is
deep and subtriangular, and is partially separated from the or-
bital by a postorbital wall.
A maxillary fragment retains a single tooth (Fig. 3DF). The
tooth is large and reminiscent of some teeth referred to Prioco-
nodon from the Aptian-Albian Arundel Formation of Maryland
FIGURE 5. Left posterior section of skull (fused postorbital + jugal +
quadratojugal + quadrate + squamosal) of Peloroplites cedrimontanus
CEUM 26331.4: A, lateral; B, posterior; C, medial views. Abbreviations:
j/jq, jugal + quadratojugal; jh, jugal horn;ltf, lateral temporal fenestra;
o, orbit; po, postorbital; poh, postorbital horn; q, quadrate; sq, squa-
mosal. Scale in cm.
FIGURE 4. Left prefrontal + lachrymal of Peloroplites cedrimontanus CEUM 26331.2: A, lateral; B, medial into the posterior portion of the nasal
cavity; C, dorsal; D, ventral into the orbit; Eanterior; F, posterior. Right prefrontal + lachrymal of Peloroplites cedrimontanus CEUM 26331.3: G,
lateral; H, medial into the posterior portion of the nasal cavity; I, dorsal; J, ventral into the orbit; K, anterior; L, posterior views. Abbreviations:dow,
dorsal orbital wall; g, groove; lf, lachrymal fenestra; nc, nasal cavity; o, orbit; ow, orbital wall. Scale in cm.
JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 28, NO. 4, 20081092
(Carpenter and Kirkland, 1998), more so than of teeth of Ed-
montonia (see Coombs, 1990). The tooth of Peloroplites cedri-
montanus has an extensive wear facet that extends the entire face
of the crown (Fig. 3F) as is seen in ankylosaurid teeth (e.g.,
Carpenter 2004:fig. 10). The tooth has 7 denticles on the anterior
margin and 6 on the posterior margin.
Both left (Fig. 7AC) and right (Fig. 7DF) rear portions of
the mandibles are preserved, although the upper half of the right
portion is folded over. These rear portions include the articular,
angular, surangular and prearticular. Laterally, the posterior sur-
face is remodeled and gives the appearance of a coossified os-
teoderm. However, a crack on the right portion, which reveals
the internal surface, does not support such an interpretation.
Medially, the adductor fossa is large and lateral-medially wider
than in Edmontonia. In addition, the fossa is separated from the
articular by a wall (Fig. 7F); such a wall is absent in Animantarx,
Edmontonia and Sauropelta. The articular cotyle is deeper rela-
tive to the size of the bone than it is in Edmontonia and Ani-
mantarx, and may correspond to the deeper and thicker retro-
articular as well. The enlarged adductor fossa for a large adduc-
tor, thick, deep retroarticular, deep articular cotyle, and
apparently large teeth suggest a strong correlation, possibly re-
lating to a diet of tougher plants than is typically thought of for
ankylosaurs (Coombs and Maryanska, 1990).
Vertebrae
The vertebrae of Peloroplites were identified by comparison
with those of Edmontonia and Sauropelta, as well as overall large
size compared with those associated with the paratype of Cedar-
pelta from the CEM site. The total vertebral count is unknown.
The atlas has not yet been found, but it was certainly not coos-
sified to the axis as in Edmontonia (Gilmore, 1930). The axis is
nearly complete, lacking parts of the neural spine and postzyg-
apophyses (Fig. 8A, B). The fragment of neural spine shows that
it was more recumbent than in Sauropelta (Fig. 8C). The centrum
is almost as tall as it is long, in marked contrast to Sauropelta and
Edmontonia where the centrum is significantly longer than tall.
FIGURE 7. Left rear portion of mandible of Peloroplites cedrimonta-
nus CEUM 12850: A, lateral; B, medial; C, dorsal views. Right rear
portion of mandible CEUM 26336: D, lateral; E, dorsal; F, medial. Ab-
breviations:af, adductor fossa; an, angular; ar, articular; cop, coronoid
process; pra, prearticular; rp, retroarticular process; sa, surangular; w,
wall of bone. Scale in cm.
FIGURE 6. Rear portion of skull of Peloroplites cedrimontanus CEUM 26331.5: A, right lateral; B, medial; C, posterior; D, anterior; E, ventral;
F, dorsal views. Abbreviations:af, adductor fossa; bo, basioccipital; bs, basisphenoid; bt, basitubera; fo, fenestra ovalis; fm, foramen magnum; fp,
frontal + parietal; jh, jugal horn;ltf, lateral temporal fenestra; nm, nuchal margin; oc, occipital condyle; paf, proatlas facet; pop, paroccipital
process; ps, parasphenoid; pt, pterygoid; q, quadrate; soc, supraoccipital crest. Scale in cm.
CARPENTER ET AL.CEDAR MOUNTAIN ANKYLOSAURS 1093
Furthermore, the ventral side of the centrum is concave, rather
than straight as in Sauropelta. The odontoid is short, but is long
in Sauropelta. The odontoid overhangs a crescent-shaped, shal-
low articular facet for the atlas. A small, flat surface, which prob-
ably represents the prezygapophysis, is seen on the right side
near the neural canal. The diapophysis is short and located just
posterior of the midline on the centrum (Fig. 8A). This is in
contrast to Sauropelta where it occurs on the neural arch. A large
parapophysis is located anteriorly on the centrum, rather than
medially as in Sauropelta.
The post-atlas cervical vertebrae consist mostly of centra (Fig.
8DU), although one nearly complete anterior cervical is present
(Fig. 8DF). The centrum of this vertebra is short, being as long
as tall. It is also sloped, so that the posterior articular face is
lower than the anterior face, a feature described by Gilmore
(1930) in Edmontonia; none of the cervicals of Sauropelta show
this degree of sloping, nor does the cervical of Cedarpelta. The
articular face is circular, rather than heart-shaped as in Edmon-
tonia nor hexagonal or horizontally ellipsoid as in Sauropelta and
Cedarpelta (Fig. 9). The neural spine is incomplete, but enough
remains to show that it was anteroposteriorly narrow and erect,
rather than short and inclined as Cedarpelta (Fig. 9). Further-
more, the anterior margin of the spine extends down between the
prezygapophyses as a ridge. The neural arch is anteroposteriorly
short, tall and erect. As a result, the postzygapophyses do not
reach the level of the posterior face of the centrum, in marked
contrast to most nodosaurids, including Edmontonia and Sauro-
pelta; the postzygapophysis extend well past the centrum in Ce-
darpelta (Fig. 9). The prezygapophysis is short and angled up-
FIGURE 8. Vertebrae of Peloroplites cedrimontanus. Axis of P.ced-
rimontanus CEUM 12812 (A,B) compared with Sauropelta AMNH 3035
(C): A,C, lateral; B, anterior views. Anterior cervical (3?) CEUM 36714
in D, anterior; E, lateral; F, posterior. Anterior dorsal CEUM 26283 in
G, anterior; H, lateral; I, posterior views. Posterior dorsal CEUM 36701
in J, anterior; K, lateral; L, posterior. Anterior caudal CEUM 50410 in
M, anterior; N, lateral; O, posterior. Anterior mid-caudal CEUM 36696
in P, anterior; Q, lateral; R, posterior. Distal caudal CEUM 12846 in S,
anterior; T, lateral; U, posterior. Abbreviations:af, atlas facet; cr, coos-
sified caudal rib; di, diapophysis; in, intervertebral notch for nerve; nc,
neural canal; ns, neural spine; o, odontoid; pa, parapophysis; poz,
postzygapophysis; pz, prezygapophysis; r, coossified rib. Scale in cm.
FIGURE 9. Vertebrae of Cedarpelta bilbeyhallorum: Anterior cervical
(3rd?) CEUM 10396: A, anterior; B, lateral; C, posterior. Synsacrum
CEUM 12163: D, dorsal; E, ventral. Anterior caudal CEUM 10258: F,
anterior; G, lateral. Anterior mid-caudal CEUM 10412: H, anterior; I,
lateral. Mid-caudal CEUM 10404: J, anterior; K, lateral. Abbreviations:
cr, coossified caudal rib; di, diapophysis; ns, neural spine; pa, parapophy-
sis; poz, postzygapophysis; pz, prezygapophysis. Scale in cm.
JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 28, NO. 4, 20081094
wards, whereas it is extremely large in Cedarpelta (Fig. 9). The
diapophysis is moderately long and steeply angled ventroposte-
riorly, rather than horizontally as in Sauropelta. As on the axis,
the parapophysis is large and located adjacent to the anterior
margin of the centrum.
Dorsal vertebrae are represented by several vertebrae, includ-
ing both anterior (Fig. 8GI) and posterior (Fig. 8JL). The
centra are amphiplatyan and lack the nodochordal projections
seen in Edmontonia. The centrum of the anterior dorsal vertebra
is shorter than tall as in Edmontonia, whereas the centrum is as
long as or longer than tall in Sauropelta. The ventral margin of
the centrum in Peloroplites is strongly concave, and less so in
Sauropelta. The prezygapophyses are steeply angled as in Sau-
ropelta, whereas they are considerably less so in Edmontonia.
The neural spine is incomplete, but the anterior margin does
extend between the prezygapophyses. The transverse processes
are angled upwards and terminate in subtriangular diapophyses.
Ventrally, a ridge extends along the transverse process to the
parapophysis located on the neural arch. The arch is short and
the neural canal subcircular. The conjoined postzygapophyses do
not extend beyond the posterior face of the centrum.
The posterior dorsal centra are as long as they are tall, thus
resembling more closely those of Sauropelta than Edmontonia,
which are taller than long. Unfortunately, the neural spines are
incomplete, thus their height is unknown. The ribs are coossified
with the vertebrae as is typical for ankylosaurs.
The synsacrum consists of six coossified vertebrae (Fig. 10A,
B), as in Silvisaurus (Carpenter and Kirkland, 1998) of which
three are probably true sacrals. The other vertebrae include a
dorsal and two caudal vertebrae. In light of other ankylosaur
synsacrals, it seems doubtful that only a single dorsal was fused
to the sacrals, although the number of additional dorsals is un-
known. The neural spines for all the synsacral vertebrae are
missing, as are most of the neural arches. These appear to have
been lost prior to burial because of encrustation over the surface.
Ventrally a pair of ridges delineates the synsacral groove as in
Edmontonia. The sacral ribs are damaged, except for the right
second. This rib retains a portion of the acetabular facet.
Caudals are represented by vertebrae from different parts of
the tail (Fig. 8MU). The anterior caudal (Fig. 8MO) has a
centrum that is wider than tall, unlike both Sauropelta and Ed-
montonia where height is about equal to width; it is taller than
wide in Cedarpelta (Fig. 9). The centrum is proportionally longer
relative to height as in Sauropelta, rather than short as in Ed-
montonia and Cedarpelta (Fig. 9). The caudal ribs are fused to
the centrum and project laterally, rather than ventrally as in
Edmontonia or anteroventrally as in Sauropelta. Proximally
there is no dorsal process or shoulderdeveloped on the caudal
rib as there is in Edmontonia, and the distal ends of the ribs are
expanded as in both Edmontonia and Sauropelta, but not in
Cedarpelta (Fig. 9). The mid-caudal vertebra (Fig. 8P, R) is the
most complete vertebra from the tail region. This specimen
FIGURE 10. Girdle parts of Peloroplites cedrimontanus. Synsacrum CEUM 50427 and iliumCEUM 50418: A, dorsal; B, ventral. Scapula+coracoids
CEUM 11706: C, dorsal; D, lateral; E, ventral; F, medial. Pubis CEUM 11626: G, dorsal; H, lateral; I, medial. The acromion is damaged. Abbre-
viations:a, acetabulum; ac, acromion; cf, coracoid fenestra; co, coracoid; gl, glenoid; i, facet for ilium; ip, ischial peduncle; is, facet for ischium; pa,
postacetabular portion of right ilium; pop, postpubic process; pp, pubic peduncle; pr, preacetabular portion of right ilium; prp, prepubic process; sc,
scapula; sr, sacral ribs; vg, ventral groove. Scale in cm.
CARPENTER ET AL.CEDAR MOUNTAIN ANKYLOSAURS 1095
shows that the caudal neural spines were laterally expanded as in
Sauropelta and Edmontonia; it is narrow and blade-like in Ce-
darpelta (Fig. 9). The postzygapophyses completely overhang the
posterior face of the centrum, in marked contrast to both Sau-
ropelta and Edmontonia; the prezygapophyses project dorsally
more as well. The neural arch is thick and encloses a subcircular
neural canal. The centrum is circular rather than heart-shaped as
in Edmontonia or hexagonal as in Sauropelta. The caudal rib
projects laterally and slightly ventrally, although the rib on the
right side is distorted ventrally. The distal caudal is damaged, but
shows that the centrum is elongated relative to its height (Fig.
8SU), whereas the height and length are about the same in
Cedarpelta (Fig. 9). The neural arch is low and encloses a circular
neural canal.
Appendicular
The right scapula and coracoid are coossified (Fig. 10CF).
They are almost complete, except for a small portion along the
dorsal margin of the coracoid and a small portion of the scapula.
The acromion process of the scapula is damaged and may be
pathological due to avulsion of the deltoideus clavicularis (scapu-
lohumeralis anterior of Coombs, 1978). The remnant of the acro-
mion process indicates that it was not as low towards the glenoid
as in Sauropelta, but rather more similar to the condition seen in
Edmontonia and to a lesser extent Animantarx. The scapular
blade is intermediate in shape between the arcuate Edmontonia
and Animantarx and the straighter Sauropelta. The posterior
margin of the scapula is rounded (Fig. 10D). The glenoid is large
and deep as it is in most ankylosaurs. A trace of the suture
between coracoid and scapula is visible within the glenoid. The
coracoid is elongated and almost as long as the scapula. It is
pierced by the coracoid foramen which is circular on the lateral
surface and ellipsoid on the medial surface where it extends onto
the scapula as a groove.
Parts of two humeri are known (Fig. 11A, B), the left one
being more complete. The proximal end is incomplete, including
the deltopectoral crest. Nevertheless, enough of the crest re-
mains to show that the humeral shaft is elongate as in Sauropelta,
Edmontonia,Animantarx, and Gastonia. Distally, the radial and
ulnar condyles are widely separated by a deep intercondylar
notch. The radial condyle is significantly lower relative to the
ulnar condyle than seen in Sauropelta or Edmontonia.
The left radius and left and right ulna are complete (Fig. 11C
E). The radius lacks the extreme expanded ends seen in Sauro-
pelta and more closely resembles that of Edmontonia, although
more slender. The ulna is long and straight, rather than bowed as
in Sauropelta; it is short and massive in Cedarpelta and Gastonia.
The olecranon process is tall and massive, and partially over-
hangs the humeral notch; such an overhang does not occur in
Sauropelta. The radial notch on the lateral surface is practically
nonexistent due to lateral crushing.
Carpals are unknown and are probably not present in anky-
losaurs. A partial manus was found in loose association and in
proximity to some of the forelimb material. The manus has a
complete set of metacarpals (Fig. 12) making it one of the few
known specimens. Lambe (1919) and Sternberg (1921) illustrate
and described a partial manus for Panoplosaurus (I-III), Ostrom
(1970) for Sauropelta (I-V), Carpenter (1990) for Edmontonia
(I-IV) and Carpenter and Kirkland (1998) for Nodosaurus (II-
IV?). The complete metacarpal set of Peloroplites allows reas-
sessment of the identities of the metacarpals in Carpenter
(1990:fig 21.17): left metacarpal I fig. 17E; right metacarpal I fig.
17F, left metacarpal II fig. 17D, left(?) metacarpal III fig. 17C,
right metacarpal IV fig 17A, and left metacarpal IV fig 17B.
Metacarpals I-IV of Edmontonia and Sauropelta and I-III of
Panoplosaurus are more similar in length than is the case of
Peloroplites.
The proximal ends of the metacarpals are faceted and fit snug-
gly against one another (Fig. 12B). Metacarpal I is the broadest
in the manus. It is blocky and sub-rectangular, with a subtrian-
gular proximal end, and no distal condyles. Interestingly, cera-
topsians also have a similar shaped metacarpal I (see Brown,
1917). Metacarpal I of Peloroplites differs from that of Edmon-
tonia and Panoplosaurus in being proportionally shorter com-
pared to its width. The remaining metacarpals of Peloroplites are
hourglass shaped, with expanded proximal and distal ends. Meta-
carpal II is about 25% taller than metacarpal I. Its proximal end
is subtriangular and the distal condyles are weakly separated.
Metacarpal II is proportionally more slender in Panoplosaurus;
FIGURE 11. Forelimb elements of Peloroplites cedrimontanus:A,
right humerus CEUM 11704; B, left humerus CEUM 10614. Left radius
CEUM 11655: A, medial; left ulna CEUM 11347 B, in lateral; right ulna
CEUM 11708 C, in lateral. Abbreviations:dp, deltopectoral crest; hn,
humeral notch; in, intercondylar notch; op, olecranon process; rc, radial
condyle; rf, radial facet; rh, radius head; uc, ulnar condyle. Scale in cm.
FIGURE 12. Manus of Peloroplites cedrimontanus:A, extensor side; B,
proximal ends of metacarpals showing how they nest against one an-
other. I-CEUM 12193, CEUM 12192, CEUM 12218; II-CEUM 12188,
CEUM 12190; III-CEUM 12189, CEUM 12222, CEUM 12220, CEUM
12221; IV- CEUM 12187, 12219; V- CEUM 12191, 12223. B] CEUM
12193, CEUM 12188, CEUM 12189, CEUM 12187, CEUM 12191.
JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 28, NO. 4, 20081096
it is stockier in Peloroplites and Edmontonia (assuming it has
been correctly identified). Metacarpal III is the most robust and
usually the longest in the manus in all families of ankylosaurs.
The distal condyles are separate in Peloroplites and Edmontonia,
but apparently not in Sauropelta and Nodosaurus. Metacarpal IV
is about the same length as metacarpal I, although considerably
more slender. Proximally it is subpentagonal, with a slight con-
cave margin for metacarpal. Distally, the condyles are not sepa-
rate, but form a single surface. Metacarpal V is the smallest of
the series, although slightly more robust than metacarpal IV. The
distal end is rounded, with no development of separate condyles.
Few of the phalanges have been identified and their placement
is mostly estimated. They are all anteroposteriorly short (Fig.
12A), have very shallow proximal articular surfaces and poorly
developed or no distal condyles. Being so short, lateral collateral
ligament pits are absent. The longest and largest phalanx is I-1.
The distal unguals are somewhat disc-shape, being wider than
long, and rounded.
The pelvis is represented by a right ilium (Fig. 10A, B) and left
pubis (Fig. 10GI). The ilium is incomplete distally and a little
medially. The preacetabular process or blade apparently di-
verges considerably, possibly 55°in contrast to 28°in Edmonto-
nia and 39°in Sauropelta. It is possible, however, that this high
degree of divergence is an artifact of not having a complete
medial surface. The postacetabular process is broad and short as
it is in Sauropelta and Edmontonia. The lateral surface of the
blade is almost straight rather than concave as in Sauropelta and
to a lesser extent in Edmontonia. The pubis is robust compared
to that of Edmontonia and Sauropelta. The lateral face of the
pubis forms the anterior wall of the acetabulum (DiCroce and
Carpenter in prepararation). The preacetabular process is short,
straight and thick; it is proportionally longer and thinner in both
Edmontonia and Sauropelta. The postpubic process is short and
is probably not complete; it is angled posteroventrally.
The left and right femora are complete (Fig. 13AD). They are
relatively straight shafted and only slightly waisted below the
fourth trochanter. The head is set at a slight upward angle. The
crista trochanteris is low and not as prominent as in Sauropelta,
and the greater trochanter is not as rugose either. An oblique
transverse ridge is present below the greater trochanter (Fig.
13E). It is damaged on the left femur and is better seen on the
right. The fourth trochanter is moderately well developed and is
located about mid-shaft on the posteromedial side. Unlike Sau-
ropelta, the shaft is not strongly waisted or constricted below the
fourth trochanter.
The right tibia is slightly crushed towards the lateral distal end
(Fig. 13FI), but is otherwise complete. The cnemial crest is short
and rounded in profile. The tibial shaft is thick throughout its
length, not slender-waisted as in Sauropelta (the medial view of
the tibia shown by Ostrom 1970, pl. 26C, D is upside down). The
astragalus is not fused to the distal end of the tibia, which is
unusual, but not unknown, in ankylosaurs. The fibular process
extends well below the astragalar facet.
Of the hind foot, only a right metatarsal (II?) and ungual (I?)
are known (Fig. 13JM). As with most quadrupedal dinosaurs,
the metatarsal is proportionally shorter and more robust than the
metacarpals. The proximal end of the metatarsal is sloped later-
ally in anterior view, as in Sauropelta, and the distal condyles
well developed, especially posteriorly. The ungual is broad
throughout its length and is distally truncated.
Family ANKYLOSAURIDAE Brown, 1908
CEDARPELTA BILBEYHALLORUM Carpenter, Kirkland,
Burge, and Bird, 2001
HolotypeCEUM 12360 partial skull.
ParatypesDisarticulated skull: including left premaxilla
CEUM 10405; left nasal fragment CEUM 10410; right prefrontal
CEUM 10421; right lachrymal CEUM 10560; right postorbital
CEUM 10352; jugal fragment CEUM 10598; left frontal CEUM
10325; parietal CEUM 10332; right squamosal CEUM 10345;
left quadrate with attached quadrojugal CEUM 10417; right
quadratojugal CEUM 10561; braincase CEUM 10267; left sur-
angular CEUM 10270; left angular CEUM 10529. Vertebrae:
cervical centrum CEUM 11288; dorsal centra CEUM 10258,
CEUM 10409, CEUM 10442, CEUM 10360; synsacrum of 2 dor-
sals, 4 sacrals, and 1 caudal CEUM 12163; first? caudal CEUM
10258; anterior caudals CEUM 10258, CEUM 10387, CEUM
10366; mid-caudals CEUM 10255, CEUM 10257, CEUM 10260,
CEUM 10261, CEUM 10262, CEUM 10349, CEUM 10400,
CEUM 10412; posterior caudals CEUM 10404, CEUM 10407.
Appendicular: right partial humerus CEUM 10258; left ulna:
CEUM 10425; left ischium, CEUM 10266; partial right ischium
CEUM 10537; fragment of right ilium CEUM10375; cervical ribs
CEUM 10248, CEUM 10445; metacarpals CEUM, 10254,
CEUM 10356, CEUM 10430, CEUM10449, CEUM 10984; pha-
langes CEUM 10247, CEUM 9970; unguals CEUM 9922, CEUM
10253. Armor: keeled plates CEUM 10526, CEUM 10359,
CEUM 10394, CEUM 10431, CEUM 10459, CEUM 10248; com-
pressed conical plates CEUM 10359, CEUM 9960, CEUM 9962,
CEUM 10548, CEUM 10414,CEUM 10441; flat osteoderm
CEUM 10338.
FIGURE 13. Hindlimb material of Peloroplites cedrimontanus: Right
femur CEUM 11319: A, anterior; B, posterior. Left femur CEUM 11321:
C, anterior; D, posterior. E, Close-up of trochanter ridge. Right tibia
CEUM 11640: F, lateral; G, anterior; H, medial; I, proximal. Metatarsal
CEUM 35866: J, anterior; K, lateral? UngualCEUM 26324: L, dorsal; M,
lateral? Abbreviations:4th, fourth trochanter; af, astragalar facet; c,
cnemial crest; cz, crushed zone; ff, fibular facet; fh, femur head; gt,
greater trochanter; in, intercondylar notch; lc, lateral condyle; le, lateral
epicondyle; mc, medial condyle; pf, popliteal fossa; tr, trochanter ridge.
Scale in cm.
CARPENTER ET AL.CEDAR MOUNTAIN ANKYLOSAURS 1097
Holotype and Paratype LocalityCEM (Locality number
EM 419), base of the Mussentuchit Member (not top of Ruby
Ranch as given by Carpenter et al, 2001).
Referred Specimenscervical CEUM 10396; caudals CEUM
10412, CEUM 10404; coracoid CEUM 10371; humeri CEUM
10256, CEUM 11629; ischium CEUM 10266; femur CEUM
11334; tibia CEUM 11640.
Additional LocalityPrice River, 2 Quarry (Locality number
EM 372), base of the Mussentuchit Member, Cedar Mountain
Formation, Emery County, Utah. Exact locality information is
available from the Prehistoric Museum, College of Eastern Utah.
Supplemental Description
The skull parts of Cedarpelta bilbeyhallorum are described by
Carpenter and coauthors (2001) and a preliminary reconstruc-
tion of the skull is presented in Figure 14. Originally, the indi-
vidual skull parts were cataloged separately, but now have been
subsumed under a single catalog number. The description of the
skull is supplemented below with that of the postcrania. The
bones of Cedarpelta from PR-2 were separated from those of
Peloroplites by comparison with the paratype material from the
CEM site. Other bones were assigned to Cedarpelta based on
their similarity to other ankylosaurids, such as Ankylosaurus,
Euoplocephalus or Saichania, which differed from those of no-
dosaurids, such as Sauropelta and Edmontonia.
The cervical vertebrae include a complete anterior cervical
(Fig. 9AC), possibly the third based on the large size of the
prezygapophyses and position of the parapophysis low and an-
teriorly located on the centrum. The neural spine is low and
dorsoposteriorly tapered. Unfortunately the neural spines on the
cervical vertebrae of Peloroplites are too incomplete for com-
parison, although it is clear that the spine does continue as a
sharp ridge between in prezygapophyses, but not in Cedarpelta.
The cervical centrum of Cedarpelta is as long anteroposteriorly
as it is tall, and is wider than it is long. In contrast, the cervical
centrum of Peloroplites is longer than tall, and is as wide as it is
long.
The dorsal vertebrae of Cedarpelta are too damaged for de-
scription. The synsacrum of Cedarpelta is complete, save for
some of the neural spines and some of the sacral ribs (Fig. 9D,
E). Assuming that there are the usual three sacrals, then there
are four sacrodorsals and two sacrocaudals. The sacral groove
developed on the ventral side is restricted to the three sacrals.
The groove is not bounded laterally by a ridge as it is in Pelo-
roplites. The neural spines are coossified in typical ankylosaur
fashion. The anterior caudal is illustrated for the first time in
Figure 9F, G. Unlike Ankylosaurus, the chevron is not coossified
to the centrum. The centrum differs from Peloroplites in that it is
almost as tall as it is wide, whereas it is considerably wider than
tall in the latter. Furthermore, the centrum is longer relative to
its height in Peloroplites than in Cedarpelta. Although the neural
spine is missing in Peloroplites, its base shows that it was wide,
whereas it is narrow in Cedarpelta. The prezygapophysis over-
hangs the centrum in Cedarpelta, but not in Peloroplites. The
mid-caudals of both ankylosaurs are more similar than are the
anterior caudals, except for the thin neural spine of Cedarpelta
versus the wider spine in Peloroplites. Distal caudals are not yet
known, thus the degree of fusion of the distal caudals anterior to
the club (if present) is unknown.
Of the pectoral girdle, only the left coracoid has been found
(Fig. 15A). The dorsal (anterior edge when in anatomical po-
sition) is incomplete. It is plate-like and is pierced by a moder-
ately large foramen. This coracoid foramen is located far from
the scapular suture.
The humerus from PR2 is more complete (Fig. 15BE) than
the one from PR1 that was associated with the holotype skull
(Fig. 15F, G). Both show the distinctive short shaft between the
deltopectoral crest and radial condyle of ankylosaurids. The
complete humerus is proportionally short and broad, with the
deltopectoral crest situated lower than in Ankylosaurus or Euo-
plocephalus, but this may be the result of anteroposterior crush-
ing because a similar situation occurs in the crushed left humerus
of Ankylosaurus (see Carpenter 2004:fig. 16). This crushing has
also affected the deltopectoral crest, which flares laterally. The
internal tuberosity is prominently distinct, unlike the condition
in Euoplocephalus or Ankylosaurus. Distally, the radial condyle
is significantly lower than the ulnar condyle, whereas in Anky-
losaurus the condition is reversed (Carpenter 2004, fig. 16); in
Euoplocephalus the ulnar condyle is lower than the radial on the
left humerus, but are at the same relative position in the right.
FIGURE 14. Reconstruction (cast) of the skull of Cedarpelta bilbeyhallorum in A, left lateral; B, dorsal; C, anterior; D, ventral. Scale in cm.
JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 28, NO. 4, 20081098
The left ulna is complete, although laterally crushed (Fig.
15HJ). It closely resembles that of the Mongolian Talarurus as
figured by Maryanska (1977:fig. 11A
2
). The olecranon is trian-
gular in profile and is lower than in Peloroplites (compare with
Fig. 11D, E). The shaft is significantly less elongate and more
robust than Peloroplites as well. The humeral notch is less poorly
developed, and the radial notch only moderately. The two radii
are similar (Fig. 15KM), although one is distorted. The least
damaged radius, a left, is short and robust, whereas it is much
longer and proportionally narrower in the nodosaurid Peloro-
plites (compare with Fig. 11C).
The ischia are relatively straight shafted (Fig. 16), a character
frequently seen in ankylosaurids (e.g., Ankylosaurus,Saichania
and Pinacosaurus). The proximal end is expanded to form the
medial wall of the acetabulum. On the medial side is a unique
node (Fig. 16C), of unknown function, as previously noted by
Carpenter and coauthors (2001).
The femur of Cedarpelta (Fig. 17AD) has the typical short
and stocky form of ankylosaurids (except Pinacosaurus), com-
pared with the long, slender nodosaurid femur of Peloroplites.
The head is well developed compared to other ankylosaurids,
although Saichania approaches this condition. The fourth tro-
chanter is non-existent, in typical ankylosaurid condition. Dis-
tally, the medial condyle is considerably larger than the lateral
condyle (they are almost equal in Ankylosaurus) and the lateral
condyle projects lower; in Ankylosaurus,Euoplocephalus,
Saichania, and Pinacosaurus the two condyles are at the same
level. The intercondylar notch is more prominent than in Anky-
losaurus, but is present in most other ankylosaurids.
The tibia is damaged proximally and distally due, in part, to
being crushed against other bones (Fig. 17EJ). Overall, it is very
robust for its length, being more like Saichania than Euoplo-
cephalus. In lateral view the proximal end is anteroposteriorly
expanded mostly due to the cnemial crest. This expansion is
greater relative to tibial length than that seen in Euoplocephalus,
but less than seen in Saichania. The fibula is straight and nearly
parallel-sided, except distally where it flares slightly (Fig. 17K,
L). Overall, it more closely resembles that of Saichania than the
tapering fibula of Ankylosaurus and Euoplocephalus.
DISCUSSION
The Cedar Mountain Formation has proven to be unusually
rich compared with most Cretaceous formations in ankylosaur
diversity and in the numbers of specimens recovered. The nodo-
saurid Peloroplites is large, matched only in size by Sauropelta
from the Cloverly Formation. It is possible that some of the large
nodosaurid material from the Cedar Mountain Formation that
has been questionably identified as Sauropelta may actually be-
long to Peloroplites. If true, that would extend the stratigraphic
range down to the middle of the Ruby Ranch Member based on
FIGURE 15. Forelimb material of Cedarpelta bilbeyhallorum.A, cora-
coid CEUM 10371. Left humerus CEUM 11629: B, anterior; C, posterior;
D, lateral; E, medial. Right humerus CEUM 10256 in F, anterior; G,
posterior. Left ulna CEUM 11669: H, lateral; I, medial; J, anterior. Left
radius CEUM 11718 in K, lateral; L, anterior. Right radius CEUM 11631
in M, medial? Abbreviations:dp, deltopectoral crest; h, head; hn, hu-
meral notch; in, intercondylar notch; it, internal tuberosity; lsr, lateral
supracondylar ridge; op, olecranon process;rc, radial condyle; rf, radial
facet; rh, radius head; uc, ulnar condyle. Scale in cm.
FIGURE 16. Left ischium of Cedarpelta bilbeyhallorum CEUM 10266:
A, lateral; B, medial showing tuberosity (arrow); C, proximal showing
medial tuberosity (arrow). Abbreviations:ac, acetabulum; if, facet for
ilium; pf, facet for pubis. Scale in cm.
CARPENTER ET AL.CEDAR MOUNTAIN ANKYLOSAURS 1099
a specimen described by Warren and Carpenter (2004). One
specimen questionably identified as Sauropelta (Carpenter et
al.,1999), cannot be referred to either that genus or to Peloro-
plites. The specimen, from the Poison Strip Sandstone Member
of the Cedar Mountain Formation, was originally described by
Bodily (1969) as Hoplitosaurus, from the Lakota Formation of
South Dakota, based on similar compressed, triangular spines.
Such spines are characteristic of polacanthid ankylosaurs (Car-
penter, 2001), which also includes Hoplitosaurus. Therefore, the
Bodily specimen probably represents an unnamed large polacan-
thid ankylosaur rather than a nodosaurid.
As for Cedarpelta, the postcrania clearly establishes it as an
ankylosaurid (contra Vickaryous et al. 2004) based on the crite-
ria established first by Coombs (1978). Therefore, the presence
of premaxillary teeth in Cedarpelta is a plesiomorphic character
in ankylosaurids as Carpenter and coauthors (2001) originally
concluded. The shamosaurine ankylosaurs (Cedarpelta,Shamo-
saurus and Gobisaurus) now include Zhongyuansaurus luoyan-
gensis recently described by Xu and coauthors (2007) as a no-
dosaurid. The specimen includes a crushed skull, partial man-
dible, partial and complete cervical and dorsal vertebrae,
anterior caudal centra, distal end of caudal vertebrae, left hu-
merus, pubis, both ischia, and ribs. The specimen was considered
to be a nodosaurid based on the narrowness of the skull (ratio of
skull length to skull width 1.4:1) and lack of an ossified tail
club. However, the skulls of shamosaurines retain the plesiomor-
phic narrow skull as measured across the dorsal surface of the
cranium (Shamosaurus 1.3:1; Gobisaurus 1.3:1; Cedarpelta 1.8:
1); thus, the ratio of Zhongyuansaurus cannot be used to identify
it as nodosaurid. In addition, the skull of Zhongyuansaurus
shows ankylosaurid, and specifically shamosaurine features
(Vickaryous et al., 2004), including low, blocky profile, covered
lateral temporal fenestra, narrow premaxillary beak, posterolat-
erally directed paroccipital process, and crescent-shaped occipi-
tal condyle. In addition, it has the characteristic straight ischium
of ankylosaurids (Coombs, 1978), and resembles that of Cedar-
pelta (Fig. 16). The end of the tail, however, clearly lacks the
distinctive osteoderms coossified with the vertebrae into a tail
club (Xu et al., 2007:fig. 2D).
The tail club has long been considered as diagnostic of the
Ankylosauridae (e.g., Coombs, 1978; Carpenter, 2001; Vickary-
ous et al., 2004), despite the club only being known among the
ankylosaurine ankylosaurids (e.g., Euoplocephalus Coombs,
1995). The fusion of the distal caudal osteoderms into the tail
club apparently occurred late in life, at least in Pinacosaurus,
because no tail club is known for 1.5 m long juveniles (Dong et
al., 1989). If Zhongyuansaurus is accepted as a shamosaurine
ankylosaurid rather than a nodosaurid, then the tail club is an
autapomorphy of adult Ankylosaurinae. Based on this hypoth-
esis, Cedarpelta, as well as the other shamosaurines, lacked a tail
club.
In conclusion, the Cedar Mountain Formation of eastern Utah
has produced the greatest diversity of Early Cretaceous ankylo-
saurs in the world. This diversity will increase over the next few
years as specimens currently under study are described. The no-
dosaurid Peloroplites cedrimontanus and shamosaurine ankylo-
saurid Cedarpelta bilbeyhallorum are comparable in size to the
5-5.5 m long Sauropelta edwardsorum from the contemporary
Cloverly Formation of Montana. What factors drove ankylosaurs
to attain such large size at the Aptian-Albian boundary remain
unknown.
ACKNOWLEDGMENTS
We thank Ben Creissler for his suggestions of the genus and
species names. Thanks to Mike Brett-Surman (USNM), Char-
lotte Holton and Mark Norell (AMNH), John Ostrom and Lyn
Murrey (YPM) for access to specimens under their care. Thanks
for review comments by Jim Kirkland and an anonymous re-
viewer.
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CARPENTER ET AL.CEDAR MOUNTAIN ANKYLOSAURS 1101
... Taohelong differs from the Qigu ankylosaur because of a greater height-to-width ratio of its anterior caudal centra, resulting in circular articular surfaces. Moreover, the orientation of the transverse processes is different and they extend in an antero-lateral direction (Yang et al., 2013: Fig. 3); Proximal caudals of the nodosaurid Peloroplites cedrimontanus from the late Early Cretaceous of North America resemble SGP 2002/8 in having short centra, that are wider than high with a transversely extending ventral groove, as well as massive transverse processes that are round to elliptical in cross-section, longer than the centrum diameter and make a broad contact with the centrum but lack a proximo-dorsal projection (Carpenter et al., 2008). Peloroplites mainly differs in the orientation of the transverse processes, which are more laterally directed; ...
... Cedarpelta bilbeyhallorum from the late Early Cretaceous of North America has anterior caudals that are short, with a transversely oriented ventral groove and a robust transverse process, which is oval in cross section and makes a broad contact with the centrum dorsally (Carpenter et al., 2008), similar to SGP 2002/8. The main difference between Cedarpelta and the Qigu ankylosaur is the orientation of the transverse process that is more laterally directed, and the height-to-width ration of the centrum, being more round in Cedarpelta and less wide (Carpenter et al., 2008); The nodosaurid Hungarosaurus tormai from the mid Late Cretaceous of Hungary is similar to SGP 2002/8 in having short proximal caudal vertebrae with a transversely extending ventral groove and prominent ventro-laterally projecting transverse processes that are at least as long as the centrum diameter and originate at a broad contact in the dorsal half of the centrum (Ő si, 2005). ...
... Cedarpelta bilbeyhallorum from the late Early Cretaceous of North America has anterior caudals that are short, with a transversely oriented ventral groove and a robust transverse process, which is oval in cross section and makes a broad contact with the centrum dorsally (Carpenter et al., 2008), similar to SGP 2002/8. The main difference between Cedarpelta and the Qigu ankylosaur is the orientation of the transverse process that is more laterally directed, and the height-to-width ration of the centrum, being more round in Cedarpelta and less wide (Carpenter et al., 2008); The nodosaurid Hungarosaurus tormai from the mid Late Cretaceous of Hungary is similar to SGP 2002/8 in having short proximal caudal vertebrae with a transversely extending ventral groove and prominent ventro-laterally projecting transverse processes that are at least as long as the centrum diameter and originate at a broad contact in the dorsal half of the centrum (Ő si, 2005). The main difference between Hungarosaurus and SGP 2002/8 is a greater height-to-width ratio of the anterior caudals and a slight proximo-dorsal projection of their transverse processes (Ő si, 2005); The derived ankylosaurid Dyoplosaurus acutosquameus from the Late Cretaceous of North America also has short proximal caudals with massive transverse processes that are elliptical in cross-section but they differ from SGP 2002/8 in the greater height-to-width ratio of the centra as well as the antero-laterally directed orientation of the transverse processes (Arbour et al., 2009); Euoplocephalus tutus from the Late Cretaceous of North America has anterior caudal vertebrae resembling the ankylosaur from the Qigu Fm. in the elliptical form of the centrum (being wider than high) that often exhibits a notochordal prominence, as well as the antero-posterior shortness of the centrum (Coombs, 1971;Arbour et al., 2009;Arbour and Currie, 2013). ...
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The first evidence of an ankylosaur from the Late Jurassic Qigu Formation of the southern Junggar Basin (Xinjiang, northwestern China) is described, based on an isolated caudal vertebra that was discovered together with fragmentary remains of other dinosaurs, including stegosaurs, sauropods, and theropods. The caudal vertebra is characterized by the following features: (i) elliptical morphology of the centrum, being wider than high; (ii) short antero-posterior length of the centrum; (iii) pronounced transversely extending ventral groove; (iv) massive transverse process, that is longer than the centrum diameter; (v) transverse process meeting the centrum high at the dorsal half and at a relatively flat angle; (vi) transverse process making a broad contact with the neural arch without forming a proximo-dorsal projection; and (vii) notochordal prominence present in the centre of the anterior articular surface. The study specimen represents only the second record of an ankylosaur from the Jurassic of Asia – aside from the slightly older Tianchisaurus from the early Upper Jurassic Toutunhe Formation, equally from the Junggar Basin. It helps to fill a gap in our knowledge of the early evolution of these armoured dinosaurs. Additionally, this discovery highlights the potential of the southern Junggar Basin to yield a rich vertebrate fauna and thus to provide an important insight into Late Jurassic ecosystems of Central Asia.
... To incorporate additional palatal variations noted in SAMA P40536, K. ieversi, and other ankylosaurians, we added a new character (178 in Arbour and Currie, 2016;and 190 in Soto-Acuña et al., 2021) that describes the position of the choanae within the palate relative to the maxillary tooth row: choanae with their anterior margins inline or within the anterior third of the maxillary tooth row (178/190:0); choanae posteriorly situated with their anterior margins at least mid-way along the tooth row (178/190:1). Ankylosaurians were scored from the literature (Eaton Jr, 1960;Sereno and Zhimin, 1992;Lee, 1996;Godefroit et al., 1999;Carpenter et al., 2001aCarpenter et al., , 2008Carpenter et al., , 2011Vickaryous et al., 2001;Hill et al., 2003;Carpenter, 2004;Kilbourne and Carpenter, 2005;Parsons and Parsons, 2009;Arbour and Currie, 2013;Arbour et al., 2014a;Leahey et al., 2015;Kinneer et al., 2016;Arbour and Evans, 2017;Yang et al., 2017;Bourke et al., 2018;Paulina-Carabajal et al., 2018;Wiersma and Irmis, 2018;Norman, 2020;Park et al., 2020); however, we could not adequately code the outgroup taxon Huayangosaurus taibaii because the condition of its choanae is unknown. Character polarity was therefore determined from Hesperosaurus mjosi (Maidment et al., 2018) and a 3D cranial model of Stegosaurus armatus (specimen number; UMNH VPC 44, sketchfab. ...
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Australian dinosaur research has undergone a renaissance in the last 10 years, with growing knowledge of mid-Cretaceous assemblages revealing an endemic high-paleolatitude Gondwanan fauna. One of its most conspicuous members is ankylosaurs, which are rare but nonetheless occur in most Australian dinosaur-bearing formations spanning the uppermost Barremian to lower Cenomanian. Here we describe a partial ankylosaur skull from the marine Toolebuc Formation exposed near Boulia in western Queensland, Australia. This skull represents the oldest ankylosaurian material from Queensland, predating the holotype of Kunbarrasaurus ieversi, which was found in the overlying Allaru Mudstone. The ankylosaur skull is encased in a limestone concretion with the maxillary tooth rows preserved only as impressions. Synchrotron radiation X-ray tomography was used to non-destructively image and reconstruct the specimen in 3D and facilitate virtual preparation of the separate cranial bones. The reconstruction of the skull revealed the vomer, palatines, sections of the ectopterygoids and maxillae, and multiple teeth. The palate has posteriorly positioned choanae that differs from the more anterior placement seen in most other ankylosaurians, but which is shared with K. ieversi, Akainacephalus johnsoni, Cedarpelta bilbeyhallorum, Gobisaurus domoculus, and Panoplosaurus mirus. Phylogenetic analyses place the new cranial material within the recently named basal ankylosaurian clade Parankylosauria together with K. ieversi. This result, together with the anatomical similarities to the holotype of K. ieversi, permits its referral to cf. Kunbarrasaurus sp. This specimen elucidates the palatal anatomy of Australian ankylosaurs and highlights one of the most ubiquitous components of Australian mid-Cretaceous dinosaur faunas.
... version). For trackmaker identification we follow the criteria of McCrea et al. (2001) and the anatomical descriptions of ankylosaurian autopodia (Maryanska, 1977;Carpenter, 1984;Pereda Suberbiola et al., 2005;Carpenter et al., 2008;Currie et al., 2011;Senter, 2011;Sissons, 2011;Herrero et al., 2017). Terms footprint, print and track are used as synonyms in a general way . 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129 130 YCRES104810_proof ■ 15 March 2021 ■ 2/ 16 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 ...
... So, the manus of at least some ankylosaurids is asymmetrical with emphasis on the inner part and very short digits. The manual phalangeal formula in nodosaurids such as Sauropelta is 2-3-4-3-2/3, with symmetrical length of digits (Carpenter, 1984), metacarpal I is shorter than that of ankylosaurids, and the whole metacarpus is more symmetrical than that of ankylosaurids (Ostrom, 1970;Carpenter, 1984;Carpenter et al., 2008;Sissons, 2011). Considering these characters, Nodosauridae features link better with Tetrapodosaurus than those of Ankylosauridae (in agreement with Carpenter, 1984 andHerrero et al., 2017). ...
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... Both of these characters are absent in theropod and sauropodomorph dinosaurs, non-thyreophoran ornithischians and basally branching representatives of this clade, such as Scutellosaurus (Colbert, 1981) and Scelidosaurus (Norman, 2019). Although some derived ankylosaurs show a similar development of the radial condyle (e.g., Ankylosaurus; Carpenter, 2004;Euoplocephalus;Arbour and Currie, 2013), this is not the case in most taxa, including basally branching taxa, such as Myrmoorapelta (MWC 6745), Gastonia (Kinneer et al., 2016), Sauropelta (Coombs, 1978), Gargoyleosaurus (Kilbourne and Carpenter, 2005), or Cedarpelta (Carpenter et al., 2008). In stegosaurs, such an expansion is present in Huayangosaurus (Zhou, 1984), Dacentrurus (Owen, 1875;Galton, 1985;Costa and Mateus, 2019), Hesperosaurus , Kentrosaurus ( Fig. 4B; Hennig, 1925;Galton, 1982), Stegosaurus Fig. 4C; Gilmore, 1914), Loricatosaurus (Galton, 1985(Galton, , 1990, Miragaia (ML 433) and a humerus referred to Chungkingosaurus (Dong et al., 1983: fig. ...
... The oblique crest extending from the distal end of the deltopectoral crest to the medial condyle seems to have an even more restricted distribution. This character is absent in the non-eurypodan thyreophorans Scutellosaurus (Colbert, 1981) and Scelidosaurus (Norman, 2019) and most ankylosaurs (e.g., Carpenter, 2004;Carpenter et al., 2008;Arbour and Currie, 2013), including the early taxon Gastonia (Kinneer et al., 2016). A notable exception within ankylosaurs seems to be the Jurassic form Myrmoorapelta, the humerus of which shows a well-developed oblique crest connecting the deltopectoral crest with the medial condyle (MWC 6745). ...
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... Many formations worldwide include more than one ankylosaur species, but only a few formations include three or more. The Aptian−Albian Cedar Mountain Formation includes at least six ankylosaur species, although these are distributed through different members of the formation (Carpenter et al., 2008). The Campanian Dinosaur Park Formation includes the nodosaurids Edmontonia rugosidens Gilmore, 1930 and Panoplosaurus mirus (Ryan & Evans, 2005) and the ankylosaurids D. acutosquameus, E. tutus, and Sco. ...
... We demonstrate this approach by comparing two models: (i) Brownian motion, and (ii) Brownian motion with a trend; this is equivalent to the comparison between a random walk and directional evolution in the Cantius analyses. If Cope's Rule is pervasive, then the trend model should receive much more model support than Brownian motion, and its directionality parameter hunt and carrano-analyZIng PhenotyPIc evolutIon In lIneages and clades 255 FIGURE 5.-Phylogeny of ornithischian dinosaurs from the composite tree of Carrano (2006) modified according to other studies (Ford and Kirkland, 2001;Weishampel et al., 2003;Novas et al., 2004;Vickaryous et al., 2004;Averianov et al., 2006;Ryan, 2007;Carpenter et al., 2008;Maidment et al., 2008;You et al., 2008;Arbour et al., 2009;Boyd et al., 2009;Dalla Vecchia, 2009;Sues and Averianov, 2009;Butler et al., 2010), ...
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Full-text available
In this chapter we discuss methods for analyzing continuous traits, with an emphasis on those approaches that rely on explicit statistical models of evolution and incorporate genealogical information (ancestor–descendant or phylogenetic relationships). After discussing the roles of models and genealogy in evolutionary inference, we summarize the properties of commonly used models including random walks (Brownian motion), directional evolution, and stasis. These models can be used to devise null-hypothesis tests about evolutionary patterns, but it is often better to fit and compare models equally using information criteria and related approaches. We apply these methods to a published data set of dental measurements in a sequence of ancestor–descendant populations in the early primate Cantius , with the particular goal of determining the best-supported mode of evolutionary change in this lineage. We also assess a series of questions about the evolution of femoral dimensions in several clades of dinosaurs, including testing for a trend of increasing body size (Cope's Rule), testing for correlations among characters, and reconstructing ancestral states. Finally, we list briefly some additional models, approaches, and issues that arise in genealogically informed analyses of phenotypic evolution.
... This comparatively open habitat apparently allowed group formation in Gastonia. In contrast, other ankylosaurs that were discovered in the uppermost part (the Mussentuchit Member) of the Cedar Mountain Formation, such as Animantarx, Peloroplites, Cedarpelta and probably Sauropelta individuals (Carpenter et al., 2001(Carpenter et al., , 2008Kinneer et al., 2016), are known from partial, usually single individuals which may indicate solitary lifestyle. However, these remains were deposited on a broad coastal plain with a high water table. ...
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Fragmentary but associated dinosaur bones collected in 1930 from the Pine River of northeastern British Columbia are identified here as originating from an ankylosaur. The specimen represents only the second occurrence of dinosaur skeletal material from the Cenomanian Dunvegan Formation and the first from Dunvegan outcrops in the province of British Columbia. Nodosaurid ankylosaur footprints are common ichnofossils in the formation, but the skeletal material described here is too fragmentary to confidently assign to either a nodosaurid or ankylosaurid ankylosaur. The Cenomanian is a time of major terrestrial faunal transitions in North America, but many localities of this age are located in the southern United States; the discovery of skeletal fossils from the Pine River demonstrates the potential for the Dunvegan Formation to produce terrestrial vertebrate fossils that may provide important new data on this significant transitional period during the Cretaceous.
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Chapter
This chapter focuses on Ankylosauria, a monophyletic clade of quadrupedal herbivorous dinosaurs characterized by the development of parasagittal osteoderms and osseous cranial ornamentation. All twenty-one taxa are clustered into one of two main lineages, Ankylosauridae or Nodosauridae. Fossil remains of ankylosaurs are found both in marine sediments and in nonmarine strata. The distribution of ankylosaur trackways and footprints is nearly global, including Asia, Europe, North America, and South America. Most of the tracks are concentrated in coastal and floodplain deposits, representing wet, well-vegetated habitats.