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SPECIAL TOPICS:
Geology June 2010 Vol.55 No.16: 1631−1635
doi: 10.1007/s11434-009-3614-5
First ceratopsid dinosaur from China and its biogeographical
implications
XU Xing1*, WANG KeBai2, ZHAO XiJin1 & LI DunJing2
1 Key Laboratory of Evolutionary Systematics of Vertebrates, Institute of Vertebrate Paleontology & Paleoanthropology, Chinese Academy of
Sciences, Beijing 100044, China;
2 Bureau of Tourism, Zhucheng, Zhucheng 262200, China
Received July 27, 2009; accepted August 12, 2009
Ceratopsid dinosaurs represent one of the best known dinosaur groups in the Late Cretaceous, and their unquestionable fossil re-
cords are exclusively restricted to western North America. Here we report a new ceratopsid dinosaur, Sinoceratops zhuchengensis
gen. et sp. nov., from the Upper Cretaceous Wangshi Group of Zhucheng, Shandong Province, China. Cladistic analysis places
this new taxon as the only known ceratopsid from outside North America, in a basal position within the Centrosaurinae. It is con-
siderably larger than most other centrosaurines but similar in size to basal chasmosaurines. Furthermore, it is more similar to
chasmosaurines than to other centrosaurines in several features, thus blurring the distinction of the two ceratopsid subgroups. This
new find not only provides significant information on the morphological transition from non-ceratopsid to ceratopsid dinosaurs,
but also complicates the biogeography of the Ceratopsidae, and further demonstrates that fossil sampling has profound effects on
reconstructing dinosaurian biogeography.
Late Cretaceous, Wangshi Group, Ceratopsidae, Centrosaurinae, biogeography
Citation: Xu X, Wang K B, Zhao X J, et al. First ceratopsid dinosaur from China and its biogeographical implications. Chinese Sci Bull, 2010, 55: 1631−1635, doi:
10.1007/s11434-009-3614-5
Ceratopsids, or derived horned dinosaurs, are large, quad-
rupedal herbivorous ornithischian dinosaurs. They are
among the best known dinosaur groups including famous
member such as Triceratops [1,2]. Previously ceratopsid
fossils have been known only from the Late Cretaceous
deposits of the Western Interior Seaway of North America
[1]. Turanoceratops tardabilis, a taxon based on fragmen-
tary, unassociated material collected from the Upper Creta-
ceous of Uzbekistan, has been assigned to the Ceratopsidae
[3,4]. This systematic proposal has been questioned [1,5,6],
yet recent studies provide additional support for the cera-
topsid affinity of Turanoceratops tardabilis [7,8]. In the
summer of 2008, we opened a large quarry near Zangjia-
zhuang, a site located in Zhucheng, Shandong Province,
China and collected numerous bones from the Upper Creta-
ceous Wangshi Group [9]. Most bones are referable to the
*Corresponding author (xingxu@vip.sina.com)
largest known hadrosaurid Shantungosaurus as in the
nearby Longgujian site [9]. However, some cranial material
is apparently not derived from hadrosaurs but is referable to
the Ceratopsidae. Here we report a new ceratopsid taxon
based on the recovered cranial material. Given that
Turanoceratops tardabilis is probably the sister taxon to the
Ceratopsidae rather than a basal member of the group [6,10]
(also see below for a detailed analysis), the new taxon
represents the only known ceratopsid outside of North
America [7,11,12], and its discovery has significant bio-
geographical implication for the Ceratopsidae.
1 Systematic paleontology
1.1 Taxonomy
Ornithischia Seeley, 1888
1632 XU Xing, et al. Chinese Sci Bull June (2010) Vol.55 No.16
Ceratopsia Marsh, 1890
Ceratopsidae Marsh, 1888
Centrosaurinae Lambe, 1915
Sinoceratops zhuchengensis gen. et sp. nov.
1.2 Etymology
Sino (China) and ceratops (horned-face, Latinized Greek);
Zhucheng (the place that produced the specimens described
here).
1.3 Holotype
Zhucheng Dinosaur Museum (ZCDM) V0010, a partial
skull with most elements of the skull roof and partial brain-
case.
1.4 Referred specimens
ZCDM V0011, a partial skull with much of the skull roof
and most of the braincase. ZCDM V0012, partial braincase.
1.5 Type locality and horizon
Zangjiazhuang, Zhucheng, Shandong Province, China. Up-
per Cretaceous Wangshi Group [9].
1.6 Diagnosis
Large centrosaurine ceratopsid with at least ten robust,
strongly curved horn-like processes along the posterior
margin of the parietals and at least four horn-like processes
on the squamosals. It is also different from other centro-
saurine in the following features: a large accessory fenestra
anterior to the antorbital fenestra, weakly undulated external
margin of the parietals and broadly based epoccipitals.
1.7 Description and comparison
The holotype skull (Figure 1) is estimated to be 180 cm in
total length (from the snout tip to the posterior end of the
parietals along the midline), being one of the largest cen-
trosaurine skulls discovered so far [1,2].
As in many other ceratopsid specimens [1,2], the cranial
sutures are obscured and thus no preserved elements could
be precisely defined by their borders. The skull has a typical
ceratopsid profile (Figure 2), being anteriorly narrow and
posteriorly much wider in dorsal view [1]. The fused parie-
tals of the holotype are 105 cm in the maximum transverse
width. Also similar to other ceratopsids [13], the snout is
proportionally long (estimated to be about 70% of the basal
skull length). As in ceratopsids, the orbit is small (estimated
to be less than 15% of the basal skull length) and its long
axis is nearly vertically oriented, different from the propor-
tionally larger orbit with an anteroposteriorly oriented long
Figure 1 Photograph of Sinoceratops zhuchengensis holotype. ZCDM
V0010 skull in right lateral (a) and right laterodorsal (b) views; ZCDM
V0010 parietals in dorsal (c), posterior (d), and ventral (e) views. Abbrevia-
tions: bp, bump; ff, frontal fontanelle; mb, median bump; mr, midline ramus;
nb, nasal bump; nh, nasal horn; ob, orbit; pb, postorbital bump; sb, supraorbi-
tal bump. Scale bar = 8 cm for (a), (b) and 6 cm for (c), (d), and (e).
axis in most more basal ceratopsians [5]. Similar to other
ceratopsids, the infratemporal fenestra is small in size and is
located considerably ventral to the orbit, though not to the
degree seen in most other ceratopsids [1]. Similar to other
ceratopsids, a highly reduced antorbital fenestra is present
along the posterodorsal border of the maxilla. A second
fenestra, much larger in size, is located anterior to the an-
torbital fenestra, as in an unnamed basal centrosaurine, and
XU Xing, et al. Chinese Sci Bull June (2010) Vol.55 No.16 1633
Figure 2 Photograph of Sinoceratops zhuchengensis ZCDM V0011.
Skull in left lateral (a) and dorsal (b) views. Abbreviations: af, accessory
fenestra; anf, antorbital fenestra; ff, frontal fontanelle; hp, horn-like process;
int, infratemporal fenestra; jn, jugal notch; lf, lateral fenestra; ob, orbit.
Scale bar = 10 cm.
derived non-ceratopsid neoceratopsian Zuniceratops, Ba-
gaceratops, and Magnirostris [10,12,14,15].
The lacrimal is fused to the adjacent elements and thus
its borders are not clear. The bone texture indicates, how-
ever, that the lacrimal is reduced in size. The nasals are
fused to each other. A relatively small nasal horn is located
about the mid-length of the snout. It is slightly curved pos-
teriorly and transversely compressed. It is about twice as
long anteroposteriorly as wide transversely at the cross sec-
tion. In the holotype, the mostly preserved nasal horn is
about 30 cm tall and about 25 cm in basal length. One rela-
tively prominent bump is located anterolateral to the nasal
horn and a less prominent one located anterior and slightly
lateral to the nasal horn.
The jugal contributes a large portion of the cheek, but its
contact relationships to the adjacent elements are not clear
due to the obscured sutures. However, it must have a large
contact with the squamosal as in other ceratopsids [2] in-
ferred from the general morphology of the postorbital area
and at the junction area of the jugal, squamosal and the
postorbital is a large, shallow depression. The postorbital
bears no horn-like structure, but a very weak postorbital
bump is present, which is located over posterodorsal corner
of the orbit. Anterior to this bump is an even weaker su-
praorbital bump, which is located over the anterodorsal
corner of the orbit.
The fused frontals bear a frontal fontanelle along the
midline, which is located medial to the postorbital bump.
Similar to that of other centrosaurines [1], it is elongate an-
teroposteriorly and narrow transversely. Posterolateral to
the frontal fontanelle are a pair of large, deep depressions
which are fenestrated along their lateral border (lateral
fenestra), a feature variably seen in some ceratopsid speci-
mens.
The bony frill is composed of the squamosals anter-
olaterally and the parietals in the rest areas as in other cen-
trosaurines. Also similar to most other centrosaurines [1],
the relatively short squamosal has a postquadrate portion
about twice as long as the prequadrate portion. It borders
the infratemporal fenestra dorsally and posterally to exclude
the postorbital from the fenestra. As in other ceratopsids [1],
the squamosal is constricted posterior to the infratemporal
fenestra to form a jugal notch. At least two relatively
straight horn-like processes, which are oriented lateropos-
terodorsally, are present along the lateral margin of the
squamosal. Each parietal bears at least five horn-like,
curved processes along its posterior and lateral margins.
They are short, robust, and oriented posterodorsally, with
the medial ones more prominent than the lateral ones. As in
other centrosaurines [1], the horn-like processes are some-
what imbricate in arrangement. The fused parietals are
slightly indented posteriorly along the midline at the poste-
rior margin and more similar to chasmosaurines [1], the
external margin of the parietals is relatively weakly undu-
lated. Also similar to chasmosaurines [1], the epoccipitals
are broad based, with the adjacent ones contacting each
other. The posterior and particularly lateral ramus is narrow
in dorsal view compared to the midline ramus. The latter is
triangular in cross-section, bearing a relatively broad sagit-
tal crest as in other centrosaurines [1]. There are at least 10
prominent bumps along the dorsal surface of the posterior
ramus and lateral ramus of the fused parietals and one on
the posterodorsal surface at the midline. These bumps are
more prominent close to the midline than laterally. Smaller
bumps are also present on the dorsal surface of the midline
ramus. There are two large parietal fenestrae, the lateral
borders of which are oriented anterolaterally as in other
centrosaurines [1].
The braincase elements are firmly fused to the skull roof
and as in other ceratopsids [1], the braincase is located ante-
rior to the parietals. The occipital condyle is large, measur-
ing about 90 cm in diameter in ZCDM 0012.
2 Discussion
We investigated the systematic position of Sinoceratops
zhuchengensis by scoring it into a recently published dataset
for ceratopsid phylogeny [16]. We also add Turanoceratops,
a taxon claimed to be a basal ceratopsid [7], and Alberta-
ceratops, a recently described basal centrosaurine [17], into
the dataset (Table 1). The data matrix was analyzed using
the NONA (ver 2.0) software package [18] and formatting
and character exploration was performed in WinClada [19].
The analysis protocol consisted of 1000 Tree Bisection and
Regrafting tree searches followed by branch swapping. Set-
tings included collapsing unsupported branches and count-
ing all states in polymorphic codings. Other settings includ-
1634 XU Xing, et al. Chinese Sci Bull June (2010) Vol.55 No.16
Table 1 Scorings for Sinoceratops, Albertaceratops, and Turanoceratops
Taxa Scoring
Sinoceratops ???????????????1[12]010111-1??1?11111????111???100101??????????????????????????
Albertaceratops 11??????????1??1201?10101?112??11[01]1?1???111?010---1?????1??11????????????0??
Turanoceratops ????????????????0???101?0???11??????????????????????????0??111??????????????
ing the character ordering follow [16]. The analysis resulted
in 2 equally most parsimonious trees with a length of 101
steps, the strict consensus of which is shown below (Figure
3). Our analysis on this dataset places Sinoceratops zhu-
chengensis within the Centrosaurinae. Derived features
uniting Sinoceratops zhuchengensis and other centro-
saurines include short parietosquamosal frill, squamosal
much shorter than parietal, and parietal epoccipital modified
into large horns [1,17]. However, in Sinoceratops zhuchen-
gensis, the parietal margin imbrication is not evident and the
parietal epoccipital at locus 2 is posterodorsally directed
rather than medially directed as in other centrosaurines
[1,17]. These features suggest that Sinoceratops zhuchen-
gensis lies in a basal position within the Centrosaurinae as
indicated by our analysis (Figure 3).
Estimated to have a skull of 180 cm long, Sinoceratops
zhuchengensis is among the largest centrosaurines and is
much larger than other basal centrosaurines in size [17].
Comparatively, chasmosaurines are in general larger than
centrosaurines in body size and the size disparity is more
evident between the basal members of the two groups [2].
However, the discovery of Sinoceratops zhuchengensis re-
moves this size disparity. Besides, Sinoceratops zhuchen-
gensis is more similar to chasmosaurines than to other cen-
trosaurines in several features, including weakly undulated
external margin of the parietals and broadly based epoc-
Figure 3 The phylogeny and geographical distribution of derived neo-
ceratopsians. The analysis of a dataset in [16] with 3 taxa added resulted in
2 equally most parsimonious trees with a length of 101 steps (CI = 0.80
and RI = 0.88). AS, Asia; NA, North America.
cipitals, thus further blurring the distinction of the two
ceratopsid subgroups.
Sinoceratops zhuchengensis also shortens the morpho-
logical gap between non-ceratopsid and ceratopsid dino-
saurs. For example, its infratemporal fenestra is located in a
position between the relatively dorsally located one in more
basal ceratopsians and extremely ventrally located one in
other ceratopsids [1]; it has a large, accessory fenestra ante-
rior to the antorbital fenestra, as in several taxa that are most
closely related to the Ceratopsidae [14]. These features sug-
gest a more complex pattern during the transition to the
Ceratopsidae. The last feature particularly indicates that
some salient features have only a brief phase in ceratopsian
evolution.
Although Turanoceratops has been recently suggested to
be the most basal known chasmosaurine [7], our analysis
rejects this systematic hypothesis. In our analysis, Tur-
anoceratops is in a position more derived than Zuniceratops
as suggested by previous analysis [7], but is placed outside
the Ceratopsidae. It lacks several ceratopsid synapomor-
phies including presence of a nasal ornamentation and re-
duced secondary ridges on maxillary and dentary teeth,
though it has double-rooted teeth. Sinoceratops zhuchen-
gensis, suggested by our analysis, is therefore the only
known ceratopsid from outside the western North America.
The absence of ceratopsids in Asia has been an intriguing
issue in dinosaurian biogeography given that all other dino-
saur groups found in Late Cretaceous deposits of North
America are also seen in Asia, such as the Tyrannosauridae,
Ornithomimidae, Troodontidae, Alvarezsauridae, Hardro-
sauridae, Pachycephalosauria, Ankylosauridae, and most
recently the Nodosauridae [12,20,21]. A quantitative study
indicates that ceratopsians have dispersed much less than
other dinosaur groups around them [12] partially due to the
endemic nature of the Ceratopsidae, the most speciose clade
of the Ceratopsia. This has been attributed to the absence of
preferred paleoenviroments, the insufficient fossil sampling
or a combination of both these factors [12]. The discovery
of the basal centrosaurine Sinoceratops zhuchengensis sug-
gests that the latter factor might be the main reason for the
rarity of the group in Asia, though limited ceratopsid pre-
ferred paleoenviroments must have also contributed to this.
Extensive collecting has recently filled the record of several
dinosaurian groups in Asia in case of the Ceratopsidae and
Nodosauriae [20], or in North America in case of the Alva-
rezsauridae [21,22]. These discoveries provide significant
new information on the dinosaurian biogeography. Given
the ceratopsian phylogeny presented in Figure 3, the Cera-
topsidae is more likely to have originated in Asia and mi-
XU Xing, et al. Chinese Sci Bull June (2010) Vol.55 No.16 1635
grate to North America. Multiple dispersal events occurred
in the ceratopsian evolution and all these dispersals might
be from Asia to North America. Such a dispersal pattern has
also been suggested for some other dinosaur groups such as
the derived Ornithomimosauria and Tyrannosauroidea [23,
24].
We thank Chen Yanming, Zou Qingzhong, and Chen Ruxia for coordinat-
ing the project, two anonymous referees for constructive comments, Zhang
Xiayu for Figure 1(a), (b), (d) and (e), Yu Tao and He Sicai for preparing
the specimens, and Jenny Ly for editing the ms. This work was supported
by the Chinese Academy of Sciences and the Zhucheng Municipal Gov-
ernment.
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