A New Maastrichtian Species of the Centrosaurine Ceratopsid

Pachyrhinosaurus from the North Slope of Alaska

Author(s): Anthony R. Fiorillo and Ronald S. Tykoski
Source: Acta Palaeontologica Polonica, 57(3):561-573. 2012.
Published By: Institute of Paleobiology, Polish Academy of Sciences
DOI: http://dx.doi.org/10.4202/app.2011.0033
URL: http://www.bioone.org/doi/full/10.4202/app.2011.0033

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A new Maastrichtian species of the centrosaurine
ceratopsid Pachyrhinosaurus from the North Slope
of Alaska
ANTHONY R. FIORILLO and RONALD S. TYKOSKI

Fiorillo, A.R. and Tykoski, R.S. 2012. A new Maastrichtian species of the centrosaurine ceratopsid Pachyrhinosaurus
from the North Slope of Alaska. Acta Palaeontologica Polonica 57 (3): 561–573.

The Cretaceous rocks of the Prince Creek Formation contain the richest record of polar dinosaurs found anywhere in the
world. Here we describe a new species of horned dinosaur, Pachyrhinosaurus perotorum that exhibits an apomorphic
character in the frill, as well as a unique combination of other characters. Phylogenetic analysis of 16 taxa of ceratopsians
failed to resolve relationships between P. perotorum and other Pachyrhinosaurus species (P. canadensis and P. lakustai).
P. perotorum shares characters with each of the previously known species that are not present in the other, including very
large nasal and supraorbital bosses that are nearly in contact and separated only by a narrow groove as in P. canadensis,
and a rostral comb formed by the nasals and premaxillae as in P. lakustai. P. perotorum is the youngest centrosaurine
known (70–69 Ma), and the locality that produced the taxon, the Kikak−Tegoseak Quarry, is close to the highest latitude
for recovery of ceratopsid remains.

Key wo r d s: Dinosauria, Centrosaurinae, Cretaceous, Prince Creek Formation, Kikak−Tegoseak Quarry, Arctic.

Anthony R. Fiorillo [[email protected]] and Ronald S. Tykoski [[email protected]],

Perot Museum of Nature and Science, 2201 N. Field Street, Dallas, TX 75202, USA.

Received 4 April 2011, accepted 23 July 2011, available online 26 August 2011.

Introduction aged faunas at lower latitudes (Fiorillo and Gangloff 2000;

Fiorillo 2004). It includes small and large theropods, hyp−
As with nearly all fossil vertebrates, the fossil record of dino− silophodontids, hadrosaurs, pachycephalosaurs, and cerato−
saurs is biased geographically towards localities in the mid− psians (Davies 1987; Parrish et al. 1987; Clemens and Nelms
dle and lower latitudes. Understanding of dinosaurs and their 1993; Fiorillo and Gangloff 2000 2001; Gangloff et al. 2005;
ecosystems at high latitudes is much poorer. This is some− Fiorillo 2008a, b; Fiorillo et al. 2009, 2010; Gangloff and
what ironic because of the intense interest in polar ecosys− Fiorillo 2010; Erickson and Druckenmiller 2011). Addi−
tems and climates at high latitudes today. tional taxa from the Prince Creek Formation include fishes
Here we present a new pachrhinosaurian ceratopsian di− and mammals (Clemens 1994; Clemens and Nelms 1993), as
nosaur from the Prince Creek Formation of the North Slope well as a variety of invertebrates and trace fossils (Flaig et al.
in Alaska. The Prince Creek Formation ranges in age from 2011).
latest Campanian to Paleocene (summarized in Flaig et al. The diversity of pachyrhinosaurs has increased in recent
2011) and consists largely of fluviatile sediments shed from years. Pachyrhinosaurus canadensis Sternberg, 1950 was
the uplift of the Brooks Range (Fiorillo et al. 2010; Flaig et viewed as a bizarre ceratopsian when first described, being
al. 2011). This unit was deposited at latitudes similar to, or the only ceratopsian then known with enlarged nasal and
even higher than the current geographic position (Witte et al. supraorbital bosses instead of facial horns or similar pro−
1987; Lawver and Scotese 1990; Lawver et al. 2002), and it cesses (Sternberg 1950; Langston 1967, 1968, 1975). The
represents a gateway that connected the faunas and floras of discovery of Achelousaurus horneri Sampson, 1995 another
the latest Cretaceous of Asia to those of North America boss−bearing centrosaurine from the Campanian Two Medi−
(Fiorillo 2008a). cine Formation of Montana showed that pachyrhinosaurs
The Cretaceous rocks in the Prince Creek Formation con− had a greater distribution in time and space (Sampson 1995).
tain the densest known concentrations of dinosaur bones of Pachyrhinosaurus lakustai Currie et al. 2008 was erected
any high latitude rock unit in the northern or southern hemi− upon multiple individuals from the Pipestone Creek bonebed
sphere (Rich et al. 1997, 2002). Though rich in terms of fossil in the Wapiti Formation (Late Campanian) of west−central
yield, the dinosaur fauna from the Prince Creek Formation Alberta, and other P. lakustai−like specimens from higher in
shows modest taxonomic diversity compared to similarly the Wapiti Formation have yet to be described (Currie et al.

Acta Palaeontol. Pol. 57 (3): 561–573, 2012 http://dx.doi.org/10.4202/app.2011.0033

562 ACTA PALAEONTOLOGICA POLONICA 57 (3), 2012

Institutional abbreviations.—DMNH, Perot Museum of Na−

156 152 148 ture and Science (formerly the Museum of Nature and Sci−
Barrow ence), Dallas, USA; NMC, Canadian Museum of Nature, Ot−
tawa, Canada; TMP, Royal Tyrell Museum of Palaeontol−
Prudhoe ogy, Drumheller, Canada.
Bay
Teshekpuk
Lake

Kikak-Tegoseak Quarry
Ocean Point Geological setting
The Kikak−Tegoseak Quarry (Fiorillo et al. 2010) hosts a

Sagavanirktok River
Umiat
monodominant, multi−taxic bonebed within the Prince Creek
Anaktuvak River

Colville River Formation (Fig. 1). The quarry, which extends at least 90 m
ver

along the bluff overlooking the Colville River in northern

r Ri

Alaska, was discovered in the mid−1990s. The depositional

ndle
er

setting was a low−energy alluvial/coastal plain that was wet

Cha
Riv

but subject to periodic drying (Fiorillo et al. 2010). The

lik
Kil

Kikak−Tegoseak Quarry has yielded dozens of fossils from a

variety of taxa such as osteichthyan fish, the theropod dino−
saurs Dromaeosaurus albertenis, Troodon formosus, cf.
Gorgosaurus libratus, an ornithomimid, and a hadrosaur, but
ALASKA
it has also produced an abundance of skeletal remains of
Pachyrhinosaurus Sternberg, 1950 (Fiorillo et al. 2010).
Arctic Circle A 5 m × 4 m excavation in 2006 yielded several large
field jackets, many of which remain to be fully prepared.
Collectively these jackets represent approximately 2 tonnes
of fossil material. Within the jackets prepared thus far, bones
are generally in contact within a fine−grained matrix though
no bones have been found articulated. The dense concentra−
tions of bones indicate a high degree of skeletal association.
All Voorhies taphonomic groups (Behrensmeyer 1975) are
represented but a thorough bone census awaits final prepara−
tion. The bone surfaces indicate little evidence of prolonged
subaerial exposure (i.e., weathering cracks) or rounding from
fluvial abrasion. Carbonized root traces are commonly found
in the matrix and on the bone surfaces. Less common but
present are the shallow borings of dermestid beetles (Fiorillo
et al. 2010). Tooth−marked bone is rare in this assemblage
and thus far amongst the prepared material only present on
the ceratopsian remains. Based on the number of prepared
occipital condyles from this quarry, the minimum number of
Fig. 1. A. Location of the Kikak−Tegoseak Quarry on Alaska’s North Slope.
Site indicated by open star. B. Photograph of Kikak−Tegoseak Quarry during
individuals of Pachyrhinosaurus is now increased to ten.
excavation. A large field jacket still in the quarry is seen in the lower left cor− The range in size of the occipital condyle diameters remains
ner of the image. consistent with assigning all remains to Pachyrhinosaurus
(sensu Fiorillo et al. 2010; Table 1).

2008). A recently described specimen (TMP 2002.76.1)

from the Dinosaur Park Formation (dated to approximately
75 Ma) of Dinosaur Provincial Park of southern Alberta Systematic paleontology
bears similarities to A. horneri and Pachyrhinosaurus, and
may represent a new taxon (Ryan et al. 2010; Sampson and Dinosauria Owen, 1842 (sensu Padian and May 1993)
Loewen 2010). Ornithischia Seeley, 1888 (sensu Padian and May
Here we erect a new pachyrhinosaur species based upon 1993)
material from the Kikak−Tegoseak site. We present a full de−
scription, a justification for erecting a new taxon, and a
Ceratopsidae Marsh, 1888 (sensu Dodson et al. 2004)
phylogenetic analysis that compares the new taxon with Centrosaurinae Lambe, 1915 (sensu Dodson et al.
close relatives. 2004)
FIORILLO AND TYKOSKI—NEW MAASTRICHTIAN CERATOPSID FROM ALASKA 563

Fig. 2. Centrosaurine ceratopsid Pachyrhinosaurus perotorum sp. nov., paratype (DMNH 22558); skull in left lateral (A), right lateral (B), dorsal (C), ven−
tral (D), and anterior (E) views. Scale bars 10 cm.

Pachyrhinosaurus Sternberg, 1950 Type material: Holotype: DMNH 21200, posterior part of parietal includ−
ing partial median bar, posterior median emargination, and autapo−
Type species: Pachyrhinosaurus canadensis Sternberg, 1950, north side
morphic anterior−facing horns. Paratypes: DMNH 22558, partial skull
of Little Bow River, east of Carmongay, Alberta, Canada, Late Creta−
ceous, Campanian. missing parietals, squamosals, right cheek region; DMNH 21201, a short
segment of right parietal transverse bar bearing an anterior rim horn.
Pachyrhinosaurus perotorum sp. nov. Type locality: All specimens collected from the Kikak−Tegoseak Quarry
Figs. 2–5. along the Colville River, North Slope, Alaska, USA. Exact coordinates
Etymology: In recognition of members of the Perot family (Margot and on file at the Museum of Nature and Science, Dallas. The palaeolatitude
H. Ross Perot and their children), who have demonstrated a long history of this quarry places it in the ancient polar world and makes it one of the
of supporting science and science education for the public. highest latitude centrosaurine localities known.

http://dx.doi.org/10.4202/app.2011.0033
564 ACTA PALAEONTOLOGICA POLONICA 57 (3), 2012

Table 1. Maximum diameters of ceratopsian occipital condyles recov− The skull (DMNH 22558) is distorted in an oblique manner,
ered and prepared from the Kikak−Tegoseak Quarry. The incomplete such that the right side is shifted dorsally relative to the left.
condyle AK 539−V−11 is included here as a voucher for the existence The sutures between most of the cranial bones are obscured
of at least a tenth individual at this site. The means of the population of by extensive fusion, a sign of relative maturity in other
these occipital condyles and those published by Dodson (1990) are
ceratopsids. In most respects, the skull of P. perotorum
75.9 mm and 68.9 mm, respectively. The standard deviations are
closely resembles both P. canadensis and P. lakustai. There−
4.65 mm and 7.08 mm, respectively. The range in size of the Kikak−
Tegoseak sample is consistent with the variation observed across fore, we focus here only on cranial features pertinent to es−
centrosaurine taxa. tablishing phylogenetic relationships between P. perotorum
and other pachyrhinosaurs. Measurements of DMNH 22558
Specimen no. Diameter (mm) are given in Table 2. They are based on the same landmarks
DMNH 22558 78.6 used by Langston (1967, 1975) and Currie et al. (2008).
DMNH 22194 77.7 Based on these measurements (Table 2) and those of occipi−
DMNH 22195 80.8 tal condyles from the quarry (Table 1), the cranial material of
DMNH 22198 76.8 P. perotorum falls within the size range of specimens of P.
DMNH 22257 79.5 lakustai (Currie et al. 2008).
AK 539−V−01 65.1 Rostral.—The rostral bone is fused to the premaxillae, and
AK539−V−12A 74.7 the right side is missing its posteroventral portion. The rostral
AK 539−V−28 73.5 lacks an anterior point or beak−like hook (Figs. 2A, B, 3A,
AK 539−V−29 76.7 B). Instead, it has an almost continuously convex lateral pro−
AK 539−V−11 – file from dorsal to ventral contacts with the premaxillae. In
contrast, the rostral of some P. lakustai specimens is sharply
Type horizon: The specimens were recovered from lower Maastrichtian
rocks (69–70 Ma) within the Prince Creek Formation (Fiorillo et al. Table 2. Measurements (in mm) of DMNH 22558, Pachyrhinosaurus
2010; Flaig et al. 2011). This qualifies P. perotorum as the youngest perotorum paratype skull. Landmarks are those used by Langston
known centrosaurine. (1975) and Currie et al. (2008). Lowercase “e” after number indicates an
estimate, or measured to a reconstructed surface.
Diagnosis.—Pachyrhinosaurus perotorum is distinguished
from other centrosaurine ceratopsids by the presence of a pair A top of nasal boss to ventral limit of maxilla 536
of short, dorsoventrally flattened horns projecting antero− B vertical height of narial opening in lateral view 92
medially and slightly dorsally from the anterior (fenestral) C maximum vertical height of skull 536
margin of the parietal transverse bar. Potential diagnostic fea− D vertical height of nasal fossa 216e
tures of P. perotorum also include paired fossae dorsal to E top of orbit to tip of supraorbital boss 72 (left),
narial fossa; a narrow median hump on the posterior part of the 25.5 (right)
nasal boss; basal sulcus of nasal boss extends ventrally to a F rostrum to posterior edge of naris 310
level lower than the dorsal rim of narial fossa; and lack of a P2 G posterior edge of naris to anterior margin of orbit 257 (left),
horn or process on posterior margin of parietal transverse bar, 280e (right)
but this feature may vary as seen in specimens of P. lakustai H anteroposterior length of orbit 116
(Currie et al. 2008). The new taxon may also be distinguished I posterior edge of orbit to supratemporal fenestra ?
by a blunt−tipped profile to the rostrum with an upturned J antorbital length 570
premaxillary and rostral ventral border. However, this condi− K height of orbit 111e
tion is so unusual for a ceratopsian that it may prove to be an L posterior edge of naris to posterior edge of nasal 301
individual oddity rather than a diagnostic feature. P. pero− boss
torum may also be differentiated from P. lakustai by lack of M bottom of orbit to bottom of jugal ?
anterior pommel on nasal boss (pommel present in some spec− N greatest width of expansions behind nares 199
imens of P. lakustai), and nasal boss and supraorbital bosses O width across lowest flanges of premaxillae 108e
that nearly contact, separated only by narrow groove (shared P least transverse width of narial bridge 53
with P. canadensis, but bosses widely separated in P. laku− Q greatest width of narial aperture 134
stai). The new species is also differentiated from P. cana− R greatest width of nasal boss 274e
densis by the presence of a rostral comb (shared with P. S least transverse width of face between nares and 190e
lakustai, but not present in P. canadensis). orbits
Description T length of nasal boss 504

Much of the fossil material collected from the Kikak−Tego− Additional measurements relevant to Currie et al. (2008: tables 3 and 4):
seak site awaits preparation. This description is based mainly
on DMNH 22558, a partial skull missing the parietosqua− Length of skull roof (tip of nasal boss to posterior 612
edge supraorbital bosses)
mosal frill and lower right cheek area (Figs. 2, 3), and the
Skull length (rostrum to occipital condyle) 756
holotype partial parietal frill section DMNH 21200 (Fig. 4).
FIORILLO AND TYKOSKI—NEW MAASTRICHTIAN CERATOPSID FROM ALASKA 565

Fig. 3. Labelled line drawings of Pachyrhinosaurus perotorum paratype (DMNH 22558) skull in left lateral (A), right lateral (B), dorsal (C), ventral (D),
and anterior (E) views. Scale bars 10 cm.

hooked (TMP 1987.55.285, TMP 1986.55.258), and there is et al. 2008] as “the Drumheller skull”). There is no evidence
a lesser beak on specimens of P. canadensis (NMC 9485 and for pre−mortem breaking or pathology of the element. The
an un−numbered specimen at the Drumheller Valley Inter− ventral margins of the rostral have a relatively rounded,
pretive Centre informally referred to in the literature [Currie broad cross−section (Figs. 2D, 3D).

http://dx.doi.org/10.4202/app.2011.0033
566 ACTA PALAEONTOLOGICA POLONICA 57 (3), 2012

Premaxilla.—The premaxillae are fused to one another, and

most of the ventral margin of the right premaxilla has eroded foramen
away. A small part of the ventral margin of the left has been
reconstructed. The premaxillae give rise to a broad narial
septum as in other centrosaurines. A pronounced ventral an−
gle protrudes below the alveolar margin of the maxilla. Dor−
sally, the premaxillae form part of the narial bridge and con−
tribute to a rostral comb like those seen in some specimens of foramen left lateral
lateral
P. lakustai (Currie et al. 2008), but not in P. canadensis sulcus horn sulcus
(Sternberg 1950; Langston 1967, 1975). The rostral comb in− right
cludes a large, overhanging protrusion even with the anterior horn
margin of the narial fossa. A second but smaller, transversely median
oriented protrusion is posterodorsal to the first. The lightly parietal
eroded dorsal surface of the rostral comb shows a faint me− bar
dian line that marks the sutural contact between premaxillae.
The line continues posterodorsally and stops at another trans−
verse ridge that marks the anterior margin of the nasal boss.
Nasal.—The nasal boss in DMNH 22558 is very large, ex−
tending from above the anterior part of the narial fossa to
above the middle of the orbit (Figs. 2A–C, 3A–C). The boss foramen
foramen
has no anterior pommel, unlike some specimens of P. lakustai
(Currie et al. 2008). The basal sulcus (Hieronymus et al.
2009), a system of small grooves and foramina that defines the
ventrolateral margin of the boss, curves low on the lateral sur−
face of the face to a level below the dorsal rim of the narial lateral median
fossa. The lateral surfaces of the boss are marked by extensive sulcus parietal
dorsoventral grooves and ridges, which produce the palisaded bar
texture typical of Pachyrhinosaurus bosses. The sides of the
boss are otherwise relatively flat and expand dorsolaterally,
making the boss the widest part of the skull anterior to the
P3 P3
parietosquamosal frill. There is a large, sub−spherical cavity
that deeply invades the bone near the posterior end of the right P2?
side of the boss (Figs. 2B, 3B). The edges of this apparent pa−
thology are well rounded, finished bone. Within the cavity are
several osseous structures, including multi−layered, onion−
skin−like bone, the broken edges of which show they are com−
prised of coarse fibers generally oriented perpendicular to the
outside surfaces.
anterior parietal horn median parietal bar
The dorsal surface of the nasal boss is irregular and pitted
as in other Pachyrhinosaurus (Figs. 2C, 3C). In dorsal view
Fig. 4. Centrosaurine ceratopsid Pachyrhinosaurus perotorum sp. nov.,
it is widest at a point over the centre of the maxillae. The an− holotype (DMNH 21200); parietal in dorsal (A), ventral (B), and right lat−
terior part of the boss is relatively flat, and slopes antero− eral (C) views. D. Reconstruction of the posterior parietal frill based upon
ventrally to meet the premaxillae where it contributes to the DMNH 21200 and a yet to be catalogued partial parietal with a P3 horn.
rostral comb. The asymmetrical dorsolateral edges are raised Scale bars 10 cm.
above the middle of the dorsal surface, producing a concav−
ity in the center of the boss. A low median ridge in the con− P. canadensis and P. lakustai (Langston 1967, 1975; Currie
cavity probably marks the union of the two nasals. et al. 2008; Hieronymus et al. 2009).
Posteriorly, the boss bears a conspicuous median hump, The nasal boss extends posteriorly to at least over the
or dome. The hump is sub−triangular in dorsal view, narrows midpoint of the orbit. It stops abruptly at a sharp ridge backed
anteriorly, and is the highest point on the skull anterior to the on each side by narrow transverse grooves reminiscent of the
frill (Figs. 2A, C, 3A, C). It is mediolaterally narrow, with transverse tunnel described in the same place in P. cana−
lower surfaces of the boss lateral to it. The anterior half of the densis (Langston 1967, 1975). The grooves are shallow later−
hump is pitted like the rest of the boss surface, but the poste− ally between the nasal and supraorbital bosses, but then enter
rior half bears anteroposteriorly oriented ridges, or "fins" a deep triangular fossa on the midline (Figs. 2C, 3C). The
(Hieronymus et al. 2009) (Figs. 2C, 3C). This matches the groove on the left side is interrupted at one point where the
texture seen on the posterior surfaces of nasal bosses in both nasal and supraorbital bosses actually contact. The trace of
FIORILLO AND TYKOSKI—NEW MAASTRICHTIAN CERATOPSID FROM ALASKA 567

the right−side groove can be followed anteroventrally where portedly absent in P. lakustai (Sternberg 1950; Langston
it is continuous with the basal sulcus. The groove on the left 1967, 1975; Currie et al. 2008).
side may also be continuous with the basal sulcus, but poor The right supraorbital boss is almost circular in dorsal view
surface preservation makes it difficult to discern. (Figs. 2C, 3C), with clearly defined edges. The anterior edge is
Within the nasal vestibule, the size and orientation of the sharp and steep walled with a deeply recessed surface poste−
narial process (= intranarial process) is notable. The narial pro− rior to it. The posterior and posteromedial margins are raised
cess is well preserved on the right side, but partially recon− abruptly above the adjacent skull surface. The dorsal surface is
structed on the left. The process is large and robust, with a nearly planar and marked by pits and ridges, some of which
thick proximal base and triangular, flat−bottomed cross−sec− are weakly aligned anterolaterally to posteromedially. The left
tion. A lateral groove on the lower part is a continuation of boss is well defined medially and posteriorly, but has only a
faint grooves and foramina that may mark the nasal−premaxil− low anterior rim and almost indiscernable lateral and postero−
lary suture. If so, the premaxilla forms the ventral part of the lateral margins (Figs. 2C, 3C). The dorsal surface bears pits
process, as reported in TMP 2002.76.1 and Diabloceratops and ridges, but the center is much lower than that of the right
(Kirkland and DeBlieux 2010; Ryan et al. 2010). The narial boss. An unusual mass of dark−coloured, pitted material was
process angles anteromedially toward the midline, similar to found in the center of the left supraorbital boss during prepara−
the condition in P. canadensis (Figs. 2E, 3E) (Langston 1975). tion (Figs. 2C, 3C). The dark material averages 10 mm or less
Above the narial fossa and posterolateral to the rostral in thickness, and appears to follow the contours of the underly−
comb is a shallow depression best seen on the right side (Figs. ing bone ridges.
2B, 3B). The fossa is defined dorsally and posteriorly by a Parietal.—The holotype parietal (DMNH 21200) consists of
curved ridge and a series of grooves and foramina that are con− the posterior part of the median parietal bar, part of the
tinuous with the basal sulcus. The ridges from each side meet right−side transverse parietal bar, and a horncore from the left
on the midline to mark the anterior tip of the fused nasals, and side of the transverse parietal bar found in position but sepa−
together form the third horizontal protrusion that makes up the rated from the rest of the parietal by an interval of missing
rostral comb. The fossa on each side is located in the same bone (Fig. 4). The dorsal surface of the parietal is marked by
place as the “tumescent bulge” or “supranasal” boss (Lang− faint vascular grooves, while the ventral surface is nearly
ston 1975: 1582) in P. canadensis (Sternberg 1950; Langston smooth. The median bar is relatively broad with an asymmet−
1967, 1975). There is no “fist−sized knob” (Sternberg 1950: rical cross−section. The right side of the median bar is dor−
111) posterodorsal to the narial fossa. No supranasal boss or sally excavated by a broad fossa where it curves to become
fossa is present in P. lakustai (Currie et al. 2008). the transverse bar. The fossa overlaps the midline. There is
Supraorbital bosses.—The supraorbital bosses of DMNH no such fossa on the left side of the median bar.
22558 are large and asymmetrical. Both are highest along the We follow the convention of Sampson (1995) when refer−
medial and posteromedial edges, and slope anterolaterally. encing parietal horns and processes. The posterior edge of
They are separated from one another by a midline groove that the parietal has only a very shallow median emargination.
rises from the same deep median fossa as the grooves that There is no P1 horn or enlargement, a character shared with
separate the nasal and supraorbital bosses. The T−shaped pat− other pachyrhinosaurs. There is little development of a pro−
tern of grooves and ridges created by the transverse and me− cess at the P2 locus, in contrast to most other centrosaurines
dian grooves matches that in P. canadensis, but which are re− which bear at least rudimentary P2 spines or hooks (Sampson

lateral
sulcus

foramen foramen

lateral
sulcus

Fig. 5. Centrosaurine ceratopsid Pachyrhinosaurus perotorum sp. nov., paratype (DMNH 21201); partial parietal preserving portion of right transverse pa−
rietal bar and anterior rim horn in dorsal (A), ventral (B), and right lateral (C) views. Scale bars 5 cm.

http://dx.doi.org/10.4202/app.2011.0033
568 ACTA PALAEONTOLOGICA POLONICA 57 (3), 2012

Pachyrhinosaurus perotorum sal and ventral surfaces. It arises from the anterior rim of the
transverse bar lateral to the P2 position on the posterior rim of
Pachyrhinosaurus lakustai the parietal, curves anterodorsally and medially, and projects
1/79 dorsally only a little above the plane of the frill (Fig. 4C). This
differs from the P1 spikes and processes in Centrosaurus or
Pachyrhinosaurus canadensis
Styracosaurus, which rise from the dorsal surface of the pari−
etal nearer the midline (Sampson 1995; Sampson et al. 1997).
3/87 TMP 2002.76.1 We refrain from applying or modifying the convention of
Sampson (1995) for describing these processes because of the
unique position of this horn in P. perotorum.
2/66 Achelousaurus horneri
A large foramen penetrates the dorsal surface of the ante−
rior parietal rim horn of Pachyrhinosaurus perotorum near
Einiosaurus procurvicornis the base. The foramen leads to a canal that passes through the
2/79
horn core and exits through a more anteromedially posi−
Rubeosaurus ovatus
tioned foramen on the ventral surface (Fig. 4A, B). Smooth
3/86
sulci issue from both foramina, suggesting a large vascular
element passed through the horn in life. The dorsolateral
Centrosaurus apertus edge of the horn is marked by a wider longitudinal sulcus
1
broader than the other grooves on the surface, which results
Styracosaurus albertensis in a weak lateral keel for the horn.
3/92
1/62
The left horn is much smaller than the right, has fewer
Centrosaurus brinkmani longitudinal grooves, but retains a broad lateral sulcus like
that on the right horn. Its base is also penetrated by large fo−
5/97 ramina on the dorsal and ventral surfaces in the same relative
Albertaceratops nesmoi positions as those on the right side horn. Asymmetry of pari−
etal ornamentation in the same individual was evidently not
4/98 Diabloceratops eatoni uncommon in centrosaurines (Sampson et al. 1997; Ryan et
al. 2001; Currie et al. 2008).
Another specimen (DMNH 21201) is identified as a short
Chasmosaurus belli
section of a second right side parietal transverse bar that also
6/95
bears an anterior rim parietal horn (Fig. 5). It shares the pres−
Triceratops ence of foramina on the dorsal and ventral sides of the horn in
Zuniceratops christopheri
the same locations as in the holotype. The horn on DMNH
21201 also has a broad sulcus along the dorsolateral edge, al−
Protoceratops
though it is smaller and restricted to the proximal end of the
Fig. 6. Results of phylogenetic analysis. Strict consensus of the three horn. The axis of the horn does not angle as far dorsally as in
equally most parsimonious trees generated by our analysis (length = 107, the holotype, and projects almost directly anteriorly and medi−
C.I. (consistency index) = 0.8131, R.I. (retention index) = 0.8291). Num− ally. The presence of at least two individuals from the Kikak−
bers at nodes are Bremer support values / Bootstrap percentages recovered Tegoseak quarry that bear anterior parietal horns is strong evi−
from 1000 replicates. dence against either specimen being from an aberrant individ−
ual. Instead, it suggests that these specimens preserve a taxo−
1995; Currie et al. 2008; McDonald and Horner 2010). This nomically diagnostic frill autapomorphy that differentiates
resembles some parietals referred to P. lakustai (Currie et al. Pachyrhinosaurs perotorum from all other centrosaurines.
2008). Another incomplete parietal still under preparation Geographic and stratigraphic range.—Type locality and ho−
bears a large, triangular P3 horn that would project postero− rizon only.
laterally from a complete frill (Fig. 4D). The location and
size of this horn matches the P3 horn in other Pachyrhino−
saurus, especially P. canadensis (Langston 1975; Currie et
al. 2008). It does not exhibit the anteroventral twist described Phylogenetic analysis
in many P3 horns of P. lakustai (Currie et al. 2008).
The holotype parietal bears small but well−developed horns We performed a phylogenetic analysis to determine the rela−
on the anterior rim of transverse parietal bar (Fig. 4), which is a tionships between Pachyrhinosaurus perotorum and other
diagnostic character for Pachyrhinosaurus perotorum. This centrosaurines. We based our analysis on that of Currie et al.
anterior horn is best preserved on the right hand side of the (2008) which was derived in large part from previous works
holotype. It is dorsoventrally flattened with a narrower lateral by Sampson (1995), Dodson et al. (2004), and Ryan (2007).
margin and bears numerous longitudinal grooves on both dor− We expanded the taxonomic content of the analysis to encom−
FIORILLO AND TYKOSKI—NEW MAASTRICHTIAN CERATOPSID FROM ALASKA 569

Pachyrhinosaurus perotorum

Pachyrhinosaurus canadensis

Pachyrhinosaurus lakustai

Pachyrostra
TMP 2002.76.1

Achelousaurus horneri

Einiosaurus procurvicornis
Pachyrhinosaurini

Rubeosaurus ovatus

Centrosaurus apertus

Styracosaurus albertensis

Centrosaurus brinkmani
Centrosaurinae
Albertaceratops nesmoi

Diabloceratops eatoni
Ceratopsidae
Chasmosaurus belli

Chasmosaurinae Triceratops
Zuniceratops christopheri
Protoceratops

AGE
90 80 79 78 77 76 75 74 73 72 71 70 69 68 67 66 65 (Ma)
Sa.

Cen. Tur. Con. Campanian Maastrichtian Dan. Stage

Late Cretaceous Pal. Epoch

Fig. 7. Results of phylogenetic analysis. One of three equally most parsimonious trees (length = 107, C.I. (consistency index) = 0.8131, R.I. (retention in−
dex) = 0.8291) from our analysis set upon the Late Cretaceous timescale. Ranges of ceratopsian taxa other than Pachyrhinosaurus perotorum from
Sampson and Loewren (2010). Timescale dates derived from Walker and Geissman (2009). Solid thick bars indicate well constrained range of a taxon.
Open thick bars indicate poorly constrained or estimated range of a taxon. Open circles at nodes indicate named node−defined clades. Arcs on branches indi−
cate stem−defined lineage names. Divergence points on tree are arbitrarily drawn, and are not based on calculations of lineage divergence dates. Abbrevia−
tions: Cen., Cenomanian; Con., Coniacian; Dan., Danian; Pal., Paleocene; Sa., Santonian; Tur., Turonian.

pass recently described centrosaurine taxa and specimens that The increased number of taxa required modification of
have added to our understanding of centrosaurine diversity characters 21, 34, and 36 from the dataset of Currie et al.
and relationships (P. perotorum, TMP 2002.76.1, Diablo− (2008) in order to accommodate wider variation in these char−
ceratops eatoni, Centrosaurus brinkmani, and Rubeosaurus acters. We also added eight characters (characters 47 through
ovatus) (Ryan and Russell 2005; Ryan 2007; Currie et al. 54) applicable to putative pachyrhinosaurs in order to more
2008; Kirkland and Deblieux 2010; Ryan et al. 2010; McDon− thoroughly test ingroup relationships. The entire character
ald and Horner 2010). This brought the number of taxa to 16, list is given in the Supplementary Online Material (SOM:
with Protoceratops andrewsi and Zuniceratops christopheri http://app.pan.pl/SOM/app57−Fiorillo_Tykoski_SOM.pdf),
designated as outgroups. The added taxa were coded from the as is the taxon−character matrix. All multistate characters were
literature. treated as unordered. Two characters were polymorphic in two

http://dx.doi.org/10.4202/app.2011.0033
570 ACTA PALAEONTOLOGICA POLONICA 57 (3), 2012

taxa, and these were treated as polymorphisms and not as un− (54% Bootstrap support) united by having nasal and supra−
certainties. Non−applicable characters were coded as missing orbital bosses that are nearly in contact, separated by only a
data rather than given the discrete character state “N” as done very narrow groove. In contrast, the nasal and supraorbital
by Currie et al (2008). The taxon−character matrix was created bosses in P. lakustai are separated by a relatively wide, smooth
and manipulated in MacClade 4.08 (Maddison and Maddison trough and by an even wider expanse in TMP 2002.76.1 (Cur−
2005), and a branch and bound parsimony analysis run using rie et al. 2008; Ryan et al. 2010). The clade was ambiguously
PAUP 4.0b10 (Swofford 2002). Bootstrap (1000 replicates) diagnosed by loss of an anteriorly or anterodorsally protruding
and Bremer support values were also calculated for the result− “pommel” on the nasal boss. There is a distinct pommel on the
ing trees. nasal bosses of both TMP 2002.76.1 and some specimens of
P. lakustai (Currie et al. 2008; Ryan et al. 2010). It should be
Results of analysis.—The analysis produced three equally noted that an equally parsimonious hypothesis from our analy−
most parsimonious trees (MPTs) with a length of 107 evolu− sis recovered P. perotorum and P. lakustai as sister taxa,
tionary transformations, a consistency index (C.I.) of 0.8131, united by the presence of a rostral comb.
and a retention index (R.I.) of 0.8291. The three trees differ
in the relative position of Pachyrhinosaurus lakustai which
is determined either as the sister taxon to P. perotorum, or to Discussion
TMP 2002.76.1, or to a clade comprising P. canadensis and
P. perotorum. A strict consensus of the three MPTs is given Phylogeny and systematics.—Our analysis was unable to
in Fig. 6. One of the three MPTs is shown in Fig. 7, set on a fully resolve relationships between P. perotorum and previ−
geological timescale and showing the temporal distribution ously described species of Pachyrhinosaurus (Fig. 6). Some
of centrosaurine taxa as listed by Sampson and Loewren differences between the three taxa include a rostral comb in
(2010). A list of apomorphies diagnosing the clades in the P. perotorum (a feature absent in P. canadensis but shared
tree shown in Fig. 7 is provided in the SOM. Relationships with P. lakustai), and nasal and supraorbital bosses that are
and character distributions discussed below are also based on nearly in contact with each other (a condition shared with P.
the cladogram in Fig. 7. canadensis, but not by P. lakustai) (Fig. 8). The nasal boss of
Centrosaurine monophyly was recovered, with Diablo− P. perotorum is also relatively deeper dorsoventrally than in
ceratops eatoni the most basal taxon of the lineage, followed the other taxa, as indicated by the basal sulcus of the boss that
by Albertaceratops nesmoi. A weakly supported clade con− extends ventrally on the face to a level lower than the dorsal
sisting of Centrosaurus apertus, Centrosaurus brinkmani, rim of the narial fossa (Figs. 2, 3, 8). The large P3 processes
and Styracosaurus albertensis was found to be on the sister of the parietals resemble those of P. canadensis in that the
lineage to the branch containing Einiosaurus procurvicornis, horns curve mostly laterally (Langston 1975), and do not ex−
Rubeosaurus ovatus, and the nasal boss−bearing pachyrhino− hibit the anteroventral twist present in some P. lakustai spec−
saurs, a result consistent with some prior analyses (Sampson imens (Currie et al. 2008).
1995; Dodson et al. 2004; McDonald and Horner 2010). A diagnostic character of P. perotorum is the presence of
Einiosaurus procurvicornis and Rubeosaurus ovatus for− anteromedially directed horns projecting into and over the
med a clade at the base of the pachyrhinosaur stem, a result parietal fenestrae from the anterior margin of the transverse
consistent with McDonald and Horner (2010). Achelousaurus parietal bar (Figs. 4, 8). The latter is especially important in
horneri was recovered as the most basal of the nasal boss− justifying the establishment of P. perotorum as distinct from
bearing taxa, a clade united by the presence of enlarged nasal other pachyrhinosaurs, because of the importance of frill or−
ornamentation covering the rostrum, and transformation of the namentation in distinguishing ceratopsid taxa. The anterior
nasal and supraorbital ornamentation into bosses in adults. edge parietal horn in P. perotorum is not only unique among
There was weak support for placement of the un−named taxon pachyrhinosaurs, but is also unknown among other cerato−
represented by TMP 2002.76.1 as either the sister to the three psids generally.
Pachyrhinosaurus species, or for P. lakustai and TMP The diversity of the centrosaurine lineage leading to
2002.76.1 together forming a clade that is the sister group to a Pachyrhinosaurus to the exclusion of Centrosaurus and its
P. canadensis + P. perotorum clade. TMP 2002.76.1 was closest kin has expanded in recent times. Sternberg (1950)
united with the Pachyrhinosaurus taxa by reduction of the considered P. canadensis so unlike other horned dinosaurs
antorbital fenestra to a small foramen, or its outright loss. that he erected a separate family for it, Pachyrhinosauridae.
The three Pachyrhinosaurus species were united by the The family−rank name was later dropped when it was demon−
presence of a nasal boss that reaches posteriorly to a point over strated that P. canadensis was allied with the “short−frilled”
the orbit. The clade was ambiguously diagnosed by pre− centrosaurines (Langston 1968, 1975). Recent cladistic anal−
maxillae bearing a rostral comb, adult nasal ornamentation yses, including ours, demonstrate there is a basal divergence
that is the mediolaterally widest part of the rostrum, the pres− of the centrosaurine lineage between those taxa more closely
ence of large, medially curled P2 hooks (uncertain in P. pero− related to Centrosaurus and those closer to Pachyrhino−
torum), and laterally or anterolaterally directed P3 horns. P. saurus (Sampson 1995; Dodson et al. 2004; McDonald and
canadensis and P. perotorum were found to be sister taxa Horner 2010). The recognition of this divergence, and the in−
FIORILLO AND TYKOSKI—NEW MAASTRICHTIAN CERATOPSID FROM ALASKA 571

Fig. 8. Comparison of line−drawing reconstructions of skulls of Pachyrhinosaurus perotorum (A), Pachyrhinosaurus canadensis (B), and Pachy−
rhinosaurus lakustai (C) in left lateral and dorsal views. A. Based on DMNH 21200 and DMNH 22558. B. Based upon NMC 9485, NMC 9602 as shown in
Langston (1975), and from Sampson and Loewen (2010). C. Derived from Currie et al. (2008). Gray fill indicates hypothetical morphology not preserved in
currently known specimens of P. perotorum. Drawings not to scale.

creased number of derived centrosaurines now known, sug− sion with earlier literature. We instead propose the name
gest that it may be useful to establish some formally defined Pachyrhinosaurini, a modification of Pachyrhinosauridae, to
clade names within Centrosaurinae. be defined as the stem−lineage of all centrosaurine ceratopsids
We choose not to resurrect or re−define the name Pachy− more closely related to Pachyrhinosaurus canadensis than to
rhinosauridae Sternberg, 1950, so as to avoid potential confu− Centrosaurus apertus (Table 3, Fig. 7). Within Pachyrhino−

Table 3. Taxonomic definitions used in this work. We recognize only monophyletic clade and lineage names with explicit ancestry−based defini−
tions. Names are given with reference to the original work in which they were coined, followed by the work that first provided an explicit and widely
accepted phylogenetic definition for the name. Taxonomic names are also identified as either node−based or stem−based, and defined.

Dinosauria Owen, 1842 – Padian and May 1993 (node): the most recent common ancestor of Triceratops and Aves, and all of that ances−
tor’s descendants.
Ornithischia Seeley, 1888 – Padian and May 1993 (stem): all taxa more closely related to Triceratops than to Aves.
Ceratopsia Marsh, 1890 – Sereno 1998 (stem): all marginocephalians more closely related to Triceratops than to
Pachycephalosaurus.
Ceratopsidae Marsh, 1888 – Dodson et al. 2004 (node): the most recent common ancestor of Centrosaurus apertus and Triceratops
horridus, and all of that ancestor’s descendants.
Centrosaurinae Lamb, 1915 – Dodson et al. 2004 (stem): all ceratopsids more closely related to Centrosaurus apertus than to Triceratops
horridus.
Pachyrhinosaurini nov. (stem): all centrosaurine ceratopsids more closely related to Pachyrhinosaurus
canadensis than to Centrosaurus apertus.
Pachyrostra nov. (node): the most recent common ancestor of Achelousaurus horneri and
Pachyrhinosaurus canadensis, and all of that ancestor’s descendants.

http://dx.doi.org/10.4202/app.2011.0033
572 ACTA PALAEONTOLOGICA POLONICA 57 (3), 2012

saurini is a division between basal forms with enlarged nasal University of Alaska, Fairbanks, USA), Paul McCarthy (The University
horns and low supraorbital horns (Einiosaurus, Rubeosaurus), of Alaska, Fairbanks, USA), Peter Flaig (The University of Texas, Aus−
and more derived taxa with nasal and supraorbital bosses in tin, USA), Kent Newman (formerly Southern Methodist University, Dal−
las, Texas, USA), Roland Gangloff (University of California, Berkeley,
adults (Achelousaurus, TMP 2002.76.1, Pachyrhinosaurus). USA), Thomas Adams (San Antonio College, San Antonio, Texas,
There is relatively good support for the clade that includes the USA), Christopher Strganac (Southern Methodist University, Dallas,
boss−bearing pachyrhinosaurins (Fig. 6), so we propose the Texas, USA), and Jason Petula (Pennsylvania State University, Harris−
name Pachyrostra and define it as the node−clade consisting of burg, USA). We also acknowledge the generosity of Philip Currie (The
the most recent common ancestor of Achelousaurus horneri University of Alberta, Edmonton, Canada) in providing digital copies of
and Pachyrhinosaurus canadensis and all of that ancestor’s his character dataset that formed the basis for our phylogenetic analysis.
descendants (Table 3, Fig. 7). We also thank Peter Dodson (The University of Pennsylvania, Phila−
delphia, USA), Andrew Farke (The Raymond M. Alf Museum of Pale−
Biogeography.—For over 200 years, observations on the ontology, Claremont, California, USA), and Michael Benton (The Uni−
versity of Bristol, Bristol, UK) who reviewed this manuscript and pro−
geographic distribution of organisms have been intimately
vided valuable comments and suggestions for improvement. The Barrow
linked to ecology and evolutionary biology (Lomolino et al. Arctic Science Consortium (BASC) and CH2M Hill (formerly Veco Po−
2004). A useful concept is the ecoregion, defined by Bailey lar Resources) provided logistical support for this project. Funding was
(1998) in such a way that the processes responsible for eco− provided by the National Science Foundation Office of Polar Programs
system differentiation make biogeographic patterns a pre− (OPP 0424594). Lastly, the Arctic Management Unit of the Bureau of
dictable phenomenon. In terms of the large−scale paleonto− Land Management provided administrative support. The specimens we
logical applications of biogeography in the Cretaceous of collected under BLM permit number AA−86367.
North America, Lehman (2001) presented evidence for lati−
tudinal zonation in the pattern of dinosaur distribution, per−
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http://dx.doi.org/10.4202/app.2011.0033