ARTICLE

Memoir of the Fukui Prefectural Dinosaur Museum 15: 27–32 (2016)© by the Fukui Prefectural Dinosaur Museum

COMMENTS ON THE VALIDITY OF THE TAXONOMIC STATUS OF
PUKYONGOSAURUS (DINOSAURIA: SAUROPODA)
Jin-Young PARK
Seodaemun Museum of Natural History
51 Yeonhui-ro 32-gil, Seodaemun-gu, Seoul 03718, Republic of Korea
ABSTRACT
Associated materials of euhelopodid sauropod, Pukyongosaurus millenniumi, were reported in the year 2001 from
Gyeongsang Basin, South Korea. However, some of the originally proposed autapomorphies for Pukyongosaurus arepresent in other sauropod taxa (i.e., somphospondylus vertebrae; ventral side of cervical centrum bent with a keel, or
flat; opisthocoelous centrum of dorsal vertebrae with long and deep pleurocoels; dorsally opened chevron). Others(i.e., highly and anteriorly positioned cervical neural spine; parapophyses situated on the lateroventral edge of thecentrum, posterior to the articular condyle) are too poorly preserved beyond observation. Therefore, the proposed
diagnostic features for Pukyongosaurus and the combinations of these characters cannot be autapomorphies, andmaking the taxonomic name nomen dubium. Rather, a somphospondylus vertebrae of Pukyongosaurus indicates it is anindeterminate Titanosauriformes. Moreover, its "probable clavicle" material is re-identiﬁed as a chevron based on themorphological and anatomical proﬁles.
Key words : South Korea, Gyeongsang Supergroup, Hasandong Formation, Titanosauriformes, clavicle, chevron

INTRODUCTION

species of Sauropoda, Pukyongosaurus millenniumi. The reported
type specimen of Pukyongosaurus (PKNU-G.102–109) consistsof four partial cervical vertebrae (PKNU-G.102–105), one partial
dorsal vertebra (PKNU-G.106), one partial rib (PKNU-G.107),
one complete chevron (PKNU-G.108), and one probableclavicle (PKNU-G.109) (Fig. 1) (According to the originaldescription, there should be seven cervical vertebrae (Donget al., 2001: 44–45) in the type specimen, but only four cervicalvertebrae were included in the plates, with no further information
about the three missing materials).
However, a number of authors have considered this taxon as anomen dubium in previous reviews (Upchurch et al., 2004; Lee
and Lee, 2006). Additionally, Mannion et al. (2013) regarded
the taxon as an indeterminate titanosauriform sauropod, basedon the presence of somphospondylus internal structure within
the presacral vertebrae. But none of the authors made detailed
comments about their reasoning. This article will focus on reexamining the taxonomic status of Pukyongosaurus. Here, theauthor will review each diagnostic feature of this taxon proposedby Dong et al (2001), and evaluate the previous identificationof specimen PKNU-G.109, which was originally described as aprobable clavicle of Pukyongosaurus.

The Gyeongsang Supergroup (Hauterivian to Campanian?) of
South Korea contains a variety of fossils (Lee et al., 2001), suchas hard shelled invertebrates (e.g., Kobayashi and Suzuki, 1936;
Yun et al., 2005), ﬁshes (e.g., Yabumoto et al., 2006; Kim et al.,2014), and reptiles (e.g., Yun and Yang, 2001; Hwang et al., 2002;
Yun et al., 2004; Lee et al., 2009). Dinosaur fossils (including
birds) were also discovered from the supergroup (e.g., Lockley
et al., 1992; Lim et al., 2001; Huh et al., 2003; Kim et al., 2005;
Lee and Lee, 2007; Lee et al., 2011). Since the first discovery
of a fragmentary humerus of a sauropod in 1973, a number ofisolated dinosaur bone materials were reported (Kim, 1983; Leeet al., 1997). In 1998, the ﬁrst associated materials of a dinosaur
were excavated from the Hasandong Formation (Aptian), whichforms the lower part of the Gyeongsang Supergroup, in Galsa-ri,
Geumseong-myeon, Hadong County (Dong et al., 2001; Lee et al.,2012; Kang and Paik, 2013). All materials were discovered fromthe same horizon. Thus, Dong et al. (2001) assumed that they
were from one individual, and assigned these to a new genus andReceived July 22, 2016. Accepted December 8, 2016.
Corresponding author ― Jin-Young PARK
E-mail: stegosaur＊hanmail.net
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JIN-YOUNG PARKINSTITUTIONAL ABBREVIATIONSAMNH, American Museum of Natural History, New York, U.S.A.;
DQ, Dinosauria International (Dana Quarry collection), New
York, U.S.A.; GCP, Grupo Cultural Paleontológico de Elche,
Spain; KUVP, University of Kansas Natural History Museum,
Kansas, U.S.A.; MUCPv, Museo de Geología y Paleontología
de la Universidad Nacional del Comahue, Argentina; PKNU,
Pukyong National University, South Korea; SMA, Sauriermuseum
Aathal, Switzerland.
COMMENTS ON DIAGNOSTIC FEATURES
According to Dong et al. (2001), Pukyongosaurus is diagnosedby the following characters: (1) somphospondylus vertebrae;(2) highly and anteriorly positioned cervical neural spine; (3)parapophyses situated on the lateroventral edge of the centrum,
posterior to the articular condyle; (4) ventral side of cervicalcentrum bent with a keel, or ﬂat; (5) opisthocoelous centrum of
dorsal vertebrae with long and deep pleurocoels; (6) dorsally
opened chevron. However, the diagnostic features of this taxon
are inadequate because of the following reasons.
1. Somphospondylus vertebrae
According to Dong et al. (2001), somphospondylus vertebrae
of Pukyongosaurus are all presacral elements. This is asynapomorphy of Titanosauriformes, and this can be seen
throughout the clade. Although not common, it occurs
within certain non-titanosauriform eusauropods as well (e.g,Mamenchisaurus) (Upchurch et al., 2004).

2. Highly and anteriorly positioned cervical neural spine
Dong et al. (2001) described the cervical vertebra of
Pukyongosaurus based on PKNU-G.102. Although the anterior
and posterior portion are not preserved and the remaining medial
region is laterally compressed and shattered, a partial neural spine
can be observed in this specimen. Dong et al. (2001) stated that thepartial neural spine in PKNU-G.102 is relatively tall and shows
an inclining dorsoanterior surface which makes it steeper thanthat of Mamenchisaurus, Omeisaurus, and Euhelopus. However,
the morphology of the neural spine can differ depending on theposition of the cervical vertebra. In many cases, cervical vertebrae
situated at the posterior part of the neck show a tendency to haverelatively higher neural spines with a steeper dorsoanterior surface(e.g., Puertasaurus, Sauroposeidon, Mamenchisaurus). On theother hand, the location of PKNU-G.102 in the neck is unclear.
With only a handful of materials, the overall shape of the cervicalneural spines of Pukyongosaurus is uncertain and therefore cannot
be compared with other sauropods.
3. Parapophyses situated on the lateroventral edge of thecentrum, posterior to the articular condyle
Dong et al. (2001) mentioned that the position of theparapophyses in Pukyongosaurus is quite similar to those ofEuhelopus and Camarasaurus. But all known sauropods tend tohave parapophyses projected lateroventrally from the centrum, andclosely situated to the condyle. Moreover, the presented cervicalmaterials in Dong et al. (2001) have incomplete anterior portions,which makes it uncertain whether the position or morphology of theparapophyses is comparable to other sauropods.

FIGURE 1. Full body silhouette of Pukyongosaurus millenniumi with preserved elements of PKNU-G.102–109 highlighted in white. A–B, Position of specimen
PKNU-G.109 as A, a clavicle according to Dong et al. (2001) and B, as a proximal chevron according to this study. Height of human silhouette = 1700 mm.

COMMENTS ON THE VALIDITY OF THE TAXONOMIC STATUS OF PUKYONGOSAURUS

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4. Ventral side of cervical centrum bent with a keel, or flat
Among the four cervical vertebrae presented in the platesof Dong et al. (2001), only one specimen (PKNU-G.102) has
a nearly complete ventral surface with an arched surface inlateral view. According to Upchurch et al. (2004), the ventrally
arched surface can be generally observed in many sauropod taxa
(e.g., Apatosaurus, Barosaurus, Bonitasaura, Camarasaurus,
Haplocanthosaurus, Qiaowanlong).
5. Opisthocoelous centrum of dorsal vertebrae with long anddeep pleurocoels
Dorsal vertebrae that are opisthocoelous can be observed in
all macronarians (Upchurch, 1998; Wilson and Sereno, 1998).
The pleurocoels in Pukyongosaurus are deep, and seem to becontinuous with chambers within the body of the centrum. Thisis a feature that can be seen in Omeisaurus and neosauropods(Upchurch et al., 2004).
6. Dorsally opened chevron
Macronarian sauropods are known by having proximallyopen chevrons (Calvo and Bonaparte, 1991; Upchurch, 1995),
but in some unusual examples, like the Chinese titanosauriforms
Dongbeititan (Wang et al., 2007), Daxiatitan (You et al.,2008), Xianshanosaurus (Lü et al., 2009), the North American
camarasaurid Camarasaurus lewisi (McIntosh et al., 1996), andthe European brachiosaurid Lusotitan (Mannion et al., 2013),
the proximal rami are linked by a bridge of bone. With only one
complete chevron material of Pukyongosaurus reported fromDong et al. (2001), the author thinks that it is too soon to conclude
whether this taxon had proximally opened or closed chevrons.
For these reasons, any features presented by Dong et al.
(2001) cannot be autapomorphies for Pukyongosaurus, nor the
combination of these characteristics can be stated as diagnostic
for this taxon. The type specimen of Pukyongosaurus has
an opisthocoelous dorsal vertebrae that can be seen in all
Macronaria, and show one synapomorphy of Titanosauriformes, asomphospondylus vertebrae. Therefore it is reasonable to conclude
that the specimen should be reclassified as an indeterminate
Titanosauriformes, and since Pukyongosaurus has no diagnostic
features, the taxon should be stated as a nomen dubium.

FIGURE 2. Specimen PKNU-G.109. Scale bar = 50 mm.

EVALUATION OF THE PROBABLE CLAVICLE MATERIAL
Dong et al. (2001) identiﬁed a bone as a probable clavicle(Fig. 2) from one of the type specimen of Pukyongosaurus(PKNU-G.109): It is almost certainly a clavicle; it is tridentshaped, being rodlike at one end and two-headed at the other.
Unfortunately, the distal end is broken. It is possibly a rightclavicle and distinctive from previously known clavicle .
In a number of studies including Dong et al. (2001), rodlike
elements associated with the pectoral girdles were interpreted
as putative clavicle materials of sauropods, and many authors

FIGURE 3. Clavicle and interclavicle in sauropod dinosaurs. A,
Reconstructed chest region of a sauropod dinosaur (based on Tschopp andMateus (2013). B, left clavicle (internal view) and C, interclavicle (externalview) from Spinophorosaurus nigerensis (GCP-CV 4229) (based on Remeset al., 2009). All scale bars = 100 mm.

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JIN-YOUNG PARK

have anticipated it to have connected the scapulae and coracoids
dorsally (e.g., Dong and Tang, 1984; He et al., 1988; Zhang,
1988; Sereno et al., 1999; Harris, 2007; Remes et al., 2009).
However, a recent study by Tschopp and Mateus (2013) leads tothe conclusion that many of the materials, that were previously
known to be clavicles, were actually interclavicle. True clavicles
from sauropods are L-shaped with a concave surface internally,
and a convex surface on the other. Usually found in pairs, theelements have a D-shaped or crescent-like in the midlength
cross-section (Tschopp and Mateus, 2013) (Fig. 3B). However,
PKNU-G.109 does not ﬁt in to this general morphology, and does
not show any related anatomical features. Therefore, the probableclavicle identification for this sauropod material is erroneous.
So far, true clavicle materials from sauropods are reported
from a non-somphospondylian macronarian sauropod (SMA0009) (Schwarz et al., 2007), the holotype of Spinophorosaurus
nigerensis (GCP-CV 4229) (Remes et al., 2009), three associatedelements (one pair: DQ-SB and one single isolated material: DQTY) from diplopocid sauropods (Galiano and Albersdörf, 2010),
from Camarasaurus sp. (KUVP 129716) (Bader et al., 2009), andtwo pairs from the Howe Quarry sauropods (?Diplodocidae) (SMA
K 24-3, 6; AMNH 30789) (Tschopp and Mateus, 2013).
Moreover, PKNU-G.109 is not morphologically similar toother sauropod interclavicles, either. According to Tschopp andMateus (2013), the interclavicle in sauropods is relatively stout,
elongated, and bowed in shape with a spatulate end posteriorly,
and a bifurcate end anteriorly (Fig. 3C). Although PKNU-G.109
shows a bifurcate shape at the anterior end, it has a moreelongated and widely divided end, which is quite different fromother known interclavicle materials in sauropods. In addition,
the shaft beneath the bifurcate end is more compressed laterally
and blade-like, which is different from the cranially ﬂat surfaced
interclavicle. Therefore, PKNU-G.109 is not an interclavicle
material.
Although only the median part of the specimen is preserved,
the elongated Y-shaped profile clearly shows that this material
is a partial chevron/haemal arch. The overall morphology of the
Y-shaped chevron can be verified in many other dinosaurian
clades; two rami forming the proximal end, each ramus
constituting the haemal canal, and a distal portion terminating as alaterally compressed blade.
The preserved medial part of the specimen is 314 mm indorsoventral length, and the preserved haemal canal is about
75 mm. Because the proximal end is not fully preserved, it isunclear whether the proximal end of this material is closed by abar of bone or not. Judging from the morphological proﬁles, it iscertain that PKNU-G.109 is much more proximally placed than
PKNU-G.108, near the base of the tail.
ACKNOWLEDGMENTS
The author would like to thank Prof. Yuong-Nam Lee(Paleontological Laboratory, Seoul National University, Republicof Korea), Mr. Hang-Jae Lee (Geological Museum, Korea Instituteof Geoscience and Mineral Resources, Republic of Korea), andMr. Jung-Kyun Kim (Graduate School of Analytical Scienceand Technology, Chungnam National University, Republic ofKorea) for valuable information. Dr. Yukimitsu Tomida (National
Museum of Nature and Science, Japan) and Dr. Junchang Lü(Institute of Geology, Chinese Academy of Geological Sciences,
China) reviewed the manuscript, and the author is grateful for
their comments. The author would also like to thank Dr. Soichiro
Kawabe (Fukui Prefectural Dinosaur Museum, Japan) for
providing useful comments on the manuscript.
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: in Korean with English abstract: in Korean: in Chinese with English adstract