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Early Cretaceous birds and pterosaurs from the Sinuiju Series, and geographic extension of the Jehol Biota into the Korean Peninsula

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Ke-Qin Gao at Peking University
Ke-Qin Gao
Quanguo Li at China University of Geosciences (Beijing)
Quanguo Li
M. Wei
M. Wei
M. Wei
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J. Paleont. Soc. Korea. Vol. 25, No. 1, (2009): p. 57-61
Abstract: The Lower Cretaceous Sinuiju Series of the Jasong Supergroup (=Jasong System of Pak and
Kim, 1996) developed in the Amnok River Basin, North Korea, has yielded a potentially important verte-
brate fauna, the taxonomic components of which are closely similar to those of the Jehol Biota best
known from the Yixian and Chiufotang (=Jiufotang) formations in western Liaoning province, China
(Chang et al., 2003; Zhou et al., 2003). The fossil-bearing beds of the Sinuiju Series consist of lacustrine
fine-grained sandstones, mudstones, tuffaceous shales and andesites with a total thickness of over 2500
meters (Pak and Kim, 1996). Stratigraphically, the Sinuiju Series rests unconformably on top of
Paleo-proterozoic metamorphic rocks, and underlies the Cretaceous Taebo Supergroup and Palaeogene
strata. The age of the Sinuiju Series was thought to be Late Jurassic (Pak and Kim, 1996), but the strata
contain the characteristic Eosestheria-Ephemeropsis-Lycoptera complex of the Jehol Biota, supporting
an Early Cretaceous age of the Series (see also Lee et al., 2001). Accordingly, the Sinuiju fauna signifies
the geographic extension of the Jehol Biota from northeast China into the Korean Peninsula. Vertebrate
fossils were found at a road-cut approximately six kilometers south of the city of Sinuiju (Figure 1). The
fossil specimens already excavated from this site include Lycoptera and sturgeon fishes, anuran amphib-
ians, several birds, possible theropod dinosaurs, and pterosaurs; none of these have been described in sci-
entific publication, and the taxonomic status has remained uncertain ever since their discovery. Most of
the fossils are preserved as dorso-ventral compressions as commonly seen in the Liaoning beds, but oc-
cur in dark shales that denote a slightly different depositional environment than the Yixian and
Chiufotang formations in western Liaoning, China.
Fossil birds:
Of the fossil birds known from the Sinuiju site, probably the most famed and also the most dubious is
the “Archaeopteryx of Korea,” found from “the third bed of the series” (Pak and Kim, 1996). This
important fossil discovery was publicized in a news report in 1993 but the fossil has never been
scientifically described. The high expectation but lack of information on its anatomical details cast
enormous doubt but heightened speculation on this conceivably important fossil bird. Until now, this
Korean bird has been known from two incomplete specimens, both have wing digits preserved that show
taxonomically significant features. The first specimen, illustrated in a photo image in a 1993 newspaper
report, consists of a nearly complete wing-supporting skeleton; the second specimen is also a partial
wing skeleton, comparable in size and structure to the first one (Figure 2). Close examination of these
specimens in a recent study has come to the conclusion that the so-called “Archaeopteryx of Korea” is
actually a confuciusornithid (Li and Gao, 2007). As clearly shown in these specimens, this fossil bird is
characterized by having: a strongly widened and subquadrangular deltopectoral crest of the humerus; a
phalangeal formula 2-3-4-x-x; the major and minor metacarpals equal in length; the proximal phalanx of
EARLY CRETACEOUS BIRDS AND PTEROSAURS FROM THE
SINUIJU SERIES, AND GEOGRAPHIC EXTENSION OF THE
JEHOL BIOTA INTO THE KOREAN PENINSULA
Ke-Qin Gao1,*, Quanguo Li2, MINGRUI WEI2, HyonUk Pak3, and Insop Pak3
1School of Earth and Space Sciences, Peking University, Beijing 100871, China
2Beijing Museum of Natural History, Beijing 100050, China
3Institute of Geology, Korean Academy of Sciences, Pyongyang, Korea
*Corresponding author: 10-6275-4883, E-mail: kqgao@pku.edu.cn
58 J. Paleont. Soc. Korea. Vol. 25, No. 1, 2009
minor digit (III) extremely short; and a semilunate bone free from the major metacarpal (Figure 2). All of
these are diagnostic features of the family Confuciusornithidae (Chiappe, 2002), and their occurrence in the
Korean fossil form indicates its affiliation with this family. Phylogenetically, the Confuciusornithidae
(Confuciusornis + Changchengornis) are more derived than Archaeopteryx, and they were the first beaked
birds with the tail vertebrae fused into a true pygostyle. Confuciusornithids were previously only known
from the Lower Cretaceous Yixian and Chiufotang formations in western Liaoning province, China, but
now it is clear that the biogeographic distribution of the family extended into the Korean Peninsula, although
the lower level taxonomy (at the generic and species level) of the Korean fossil bird is still uncertain.
Other fossil birds from the Sinuiju Series are known from several specimens that give evidence of the
occurrence of enantiornithine and even more advanced ornithurine birds in the Cretaceous System of the
Korean Peninsula. One of these specimens has both left and right feet well preserved, and shows
taxonomically significant features including: three unfused metatarsals; MT III-IV equal in length; and
MT II significantly shorter than MT III-IV. The combination of these features indicates the possible
enantiornithine affiliation of this bird. Enantiornithines are a diverse group of extinct birds primitively
retaining teeth, and are commonly known from Mesozoic deposits worldwide. Another specimen represents
a large bird with a tibia approximately 5.0 cm long. The specimen shows a partly preserved wing digit
with a very large claw in the hand, and has a robust and elongated pygostyle of more than 3.5 cm in
length. Without more detailed information on its anatomy, however, the taxonomic affinity of this large
bird cannot be ascertained until more complete material becomes available.
Pterosaurs:
Besides the above-mentioned bird fossils, the best-preserved material so far collected from the Sinuiju
Series is the skeleton of a pigeon-sized pterosaur (Figure 3). The specimen is approximately 25 cm long
from the tip of the snout to the end of the tail, and probably had a wingspan of more than 80 cm. The skull
is bilaterally compressed, but the postcranial skeleton is dorso-ventrally compressed as preserved. The
skull is short and deep, and the neck is evidently shorter than the trunk. The pectoral girdle has the
Fig. 1. Map showing the geographic location of the Sinuiju (solid dot) in relation to Beipiao and other major fossil sites
(solid dots) in western Liaoning province, China.
Gao et al.
-
Early Cretaceous Birds And Pterosaurs From The Sinuiju Series & Geographic Extension 59
scapula and coracoid fused, and the rod-like coracoids meet posteromedially at an acute angle to form a
V-shaped structure. The tail, although incompletely preserved, is probably greatly reduced to a short
stump similar to the pygostyle in advanced birds. All of these characters suggest that the Korean
pterosaur can be confidently recognized as a member of the family Anurognathidae. As a family classified
within the suborder Rhamphorhynchiodea (Wellnhofer, 1978; Unwin, 2006), the Anurognathidae include
the type genus Anurognathus and Batrachognathus; both were small to medium-sized pterosaurs (wingspan
~50 cm) known from the Upper Jurassic Solnhofen lithographic limestones and the Upper Jurassic of
Karatau, Kazakhstan. More recent discovery of Jeholopterus from the Daohugou beds, Inner Mongolia,
China, has extended the stratigraphic range of the family to the Middle Jurassic as the fossil beds have
been independently dated at 164-165 Ma (Chen et al., 2004; Yang and Li, 2004; Liu et al., 2006; Gao and
Ren, 2006; contra Wang et al., 2002: Lower Cretaceous Yixian Formation); and now the new fossil
discovery from North Korea may well provide the evidence for a geologically younger range extension
of the family into the Upper Cretaceous.
Compared to Jeholopterus from China, the immediate differences that can be recognized in the
Korean form are: ten rib-bearing trunk vertebrae (vs. 12-13); synsacrum formed by fusion of at least
seven sacral vertebrae (vs. by three); scapula and coracoid roughly equal in length (vs. coracoid about
half length of the scapula); and greater elongation of the ulna/radius segment to twice the length of the
humerus. Although it can be recognized as a member in the family Anurognathidae, the taxonomy of this
Korean pterosaur at the generic and species level cannot be determined before a thorough study of the
available specimens.
Fig. 2. Representative bird fossils from Sinuiju, North Korea: A-B, wing skeleton and claws of a confuciusornithid
(specimen of the “Archaeopteryx of Korea”); C-D, partial forelimb of a confuciusornithid; E-F, feet and the claws o
f
an enantiornitine; G, a large specimen with a long pygostyle. Abbreviations used in this figure: alc, alular metacarpal;
hu, humerus; mac, major metacarpal; mic, minor metacarpal; MT, metatarsal; ph, phalanx; rad, radius; ul, ulna;
60 J. Paleont. Soc. Korea. Vol. 25, No. 1, 2009
In general, several bird fossils from the Sinuiju Series, North Korea, document the occurrence of
confuciusornithid and enantiornithine birds in the Upper Cretaceous of the Korean Peninsula, and may
include some other forms that have not been previously recognized in the fossil record. The pterosaur,
known from a single specimen, can be referred to the family Anurognathidae. Although not yet formally
Fig. 3. Part and counter-part skeleton of an anurognathid pterosaur from the Sinuiju Series.
Gao et al.
-
Early Cretaceous Birds And Pterosaurs From The Sinuiju Series & Geographic Extension 61
named and described, this fossil extends the stratigraphic range of the family into the Upper Cretaceous
in the Korean Peninsula. The Sinuiju fossils provide arguably the most significant evidence for a
geographical extension of the Jehol Biota from northeastern China into the Korean Peninsula, and
forthcoming scientific study of these specimens will undoubtedly provide a significant source of
information for a better understanding of the evolution of the Jehol Biota.
ACKNOWLDEGEMENT
We thank many people who greatly helped this research: Wei Mingrui, Wang Wenli, and Bi Haiyan
(Beijing Museum of Natural History). Thanks are extended to Alex Kellner (National Museum of Brazil)
for discussion on pterosaur taxonomy, and to Professor R. C. Fox (University of Alberta, Canada) for
reading and improving the manuscript. This research was supported by the National Natural Science
Foundation of China (NSFC 40532008), and by the Beijing Municipal Natural Science Foundation
(5982009). This subject was given a talk at the 2009 Goseong International Symposium with the support
of 2009 Goseong International Symposium Committee.
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투고일 : 2009년 4월 15일, 심사일 : 2009년 4월 16일, 심사완료일 : 2009년 4월 30일
... NJU-57003 is another long-tailed specimen from the Mutoudeng locality (Daohugou Beds, Tiaojishan Formation), only preliminarily described . A relatively complete specimen from the Early Cretaceous of North Korea also awaits description (Gao et al., 2009), as well as a fragmentary specimen comprised of wing elements from the Middle Jurassic (Aalenian/Bajocian) Bakhar deposits of Central Mongolia (Bakhurina & Unwin, 1995). Finally, the poorly-known Mesadactylus ornithosphyos, based on the holotype BYU 2024 (a synsacrum) from the Kimmeridgian-Tithonian Morrison Formation of the USA (Jensen & Padian, 1989), is a potential anurognathid (see Bennett, 2007). ...
... Despite being crushed to the point of obliterating many details, the specimen is rather complete and provides new information for the group, including the first record of an anurognathid skull exposed in mostly lateral view. In other specimens, the skull is either exposed in mostly internal view, as in the holotype of Anurognathus ammoni (Döderlein, 1923;Wellnhofer, 1975;Bennett, 2007), or dorsoventrally crushed, as in all other specimens that preserve a skull (Riabinin, 1948;Ji & Ji, 1998;Bennett, 2007;Gao et al., 2009;Lü & Hone, 2012;Lü et al., 2018). ...
... In Vesperopterylus, the deltopectoral crest of the humerus is also trapezoidal, very similar in shape to Anurognathus (see Lü et al., 2018). In the North Korea specimen, the deltopectoral crest of the humerus seems to be trapezoidal as well (Gao et al., 2009). Furthermore, even though the "alate" condition seen in Jeholopterus is unique to it, it is still very similar to the trapezoidal conditions of Anurognathus and Vesperopterylus, differing only in being longer and more curvedthey are thus all coded as "trapezoidal" in our analysis (see Supplemental Information). ...
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... The contemporaneous or subcontemporaneous biotas occur in many other areas in NE Asia, such as Hebei, Inner Mongolia, Shandong, Jilin, and Heilongjiang provinces of China, Japan, Korea, and Transbaikalia of Russia, and share with the Jehol Biota more or less typical Jehol fossils. In addition to some biostratigraphically important invertebrates like Eoestheria and Cypridea, several other vertebrate taxa have also been reported in these regions, including the fishes Acipenseriformes, Osteoglossomorphs (e.g., Lycoptera) (Fig. 2D), the aquatic or semiaquatic reptilian Choristoderes (e.g., Monjurosuchus) (Fig. 2E) (42)(43)(44), the Squamates, the dinosaurian Psittacosauridae (e.g., Psittacosaurus) (Fig. 2F), and the avians Confuciusornithidae (9,45), Enantiornithes, and Ornithuromorpha, etc. (46). Obviously, the Jehol Biota and its contemporaneous biotas have significantly expanded laterally in East and Central Asia during the middle Early Cretaceous. ...
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Thesis
  • Aug 2020
Numerous pterosaur traces have been reported from the Jangdong Formation, Hwasun Seoyuri Dinosaur Tracksite, Jeollanam-do Province, Korea. The Jangdong Formation in the Neungju Basin is well-known for abundant dinosaur trackways and study on dinosaur acceleration phase using for the trackways. In this thesis, a detailed description of pterosaurian traces, their paleoecology, and the possible tracemaker of ichnites are conducted using four new pterosaur trace-bearing slabs and two previously reported slabs. The pterosaur tracks in the formation are classified as Pteraichnus isp. and one of the smallest ones in the Late Cretaceous. The pterosaur tracks in Seoyuri are yielding with manus-dominated and randomly oriented in high density. Some tracks are discovered in the states with claw marks. These characters indicate semi-floating behavior with feeding, possibly. The concentrated tracks and variation in size could be interpreted as gregarious behavior or multi-flock of pterosaurs. The considerable tracemaker of the Pteraichnus isp. in Hwasun is very rare in the Late Cretaceous. To date, previous reported small-sized pterosaurs in the Late Cretaceous are included in ornithocheiroidea and only discovered in North America (British Colombia, and Montana) and Arabia (Lebanon), but not in East Asia. Therefore, the tracemaker of the Pteraichnus isp. in Hwasun would be not reported ornithocheiroidea, inhabited in the Late Cretaceous Korean Peninsula. This global distribution of small-sized pterosaurs could interpret as their deficiency in the store of a lineage related to the lack of the contemporary Lagerstätten in the Late Cretaceous.
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Cretaceous fossil birds from China
Article
  • Jan 2024
The explosive radiation of early birds has been well documented by the extraordinary discoveries of the Early Cretaceous fossil birds from China in the past three decades. They have greatly expanded the diversity, disparity, and temporal distribution of Mesozoic birds, refining our knowledge of the evolutionary path leading to the characteristic avian body plan, such as feathers and powered flight, locomotion and habitat differentiation, diet and digestion, reproduction and development, feather colors and display. In addition, studies on the geological background of the Jehol Biota that produced the majority of the Chinese Cretaceous birds have provided important clues to our understanding of the taphonomy as well as the interaction between deep geology process and biological evolution in northeastern China and East Asia during the Early Cretaceous. Future multidisciplinary studies are expected to advance our knowledge of how the paleogeography of Early Cretaceous birds was formed, what had impacted the changes of the diversity of early birds in China, and the roles of early birds had played in the Early Cretaceous terrestrial ecosystem.
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A new confuciusornithid (Aves: Pygostylia) from the Early Cretaceous increases the morphological disparity of the Confuciusornithidae
Article
  • Jan 2019
Confuciusornithidae is the clade of Early Cretaceous birds most rich in materials and plays a central role in our understanding of the evolution of avian horny beaks and pygostyles. A handful of specimens demonstrate that this avian group is distinguishable from other basal birds by their robust, toothless upper and lower jaws, a fused scapulocoracoid and a tiny claw on the middle manual digit, among other features. Here, we report a new taxon of Confuciusornithidae, Yangavis confucii gen. et sp. nov., from the Early Cretaceous Jehol Biota, northeastern China. This new bird, however, has a normal-sized major digit claw, as in other basal birds, which was probably regained independently as Confuciusornithidae evolved, based on our phylogenetic study. Unfortunately, the biological significance of this trait is unclear owing to a lack of analogues in modern birds (manual claws are completely lost in adults). Yangavis confucii is differentiated from other confuciusornithids by its proportionally much longer forelimb. Our morphometric analysis indicates that the morphospace of Confuciusornithidae, with the addition of Y. confucii, is greatly broadened to a degree that it overlaps with the Early Cretaceous Ornithuromorpha and Enantiornithines, indicating that the biological diversity of confuciusornithids is greater than previously thought.
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Phylogenetic relationships among basal birds
Conference Paper
Full-text available
  • Jan 2001
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A nearly completely articulated rhamphorhynchoid pterosaur with exceptionally well-preserved wing membranes and ?hairs? from Inner Mongolia, northeast China
Article
Full-text available
  • Feb 2002
We report a new and nearly completely articulated rhamphorhynchoid pterosaur, Jeholopterus ningchengensis gen. et sp. nov., with excellently preserved fibres in the wing membrane and “hairs” in the neck, body and tail regions. Many of its characteristics such as a short neck, short metacarpals and distinctively long fifth pedal digit are characteristic of rhamphorhynchoids. The new species can be further referred to the ‘strange’ short-tailed rhamphorhynchoid family Anurognathidae. It is much more complete than the other known members of the family, namely, Anurognathus from Solnhofen, Germany, Batrachognathus from Karatau, Kazakhstan, and Dendrorhynchoides from Beipiao, Liaoning Province, China. The new pterosaur also shows that the wing membrane is attached to the ankle of the hind limb. The pedal digits are webbed. Furthermore, the “hair” of Jeholopterus bears some resemblance to the hair-like integumental structures of the feathered dinosaur Sinosauropteryx although there is yet no direct evidence to argue for or against their homology.
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UPb zircon age for the Daohugou Biota at Ningcheng of Inner Mongolia and comments on related issues
Article
  • Nov 2006
SHRIMP U-Pb zircon dating was carried out for the Daohugou Biota near Ningcheng of Inner Mongolia and for lavas overlying or underlying salamander-bearing strata at Reshuitang in Lingyuan of West Liaoning. The results suggest that the Daohugou Biota occurred at an interval from 168 Ma to 164–152 Ma. Both the Daohugou Biota and the salamander-bearing fossil assemblage are the same biota and thus developed from 168 to 152 Ma, i.e. from late Middle Jurassic to the early Late Jurassic. The Daohugou Biota-bearing rocks, resting on the Jiulongshan Formation in disconformity and being overlain in unconformity by Late Jurassic Tuchengzi Formation and Early Cretaceous rocks containing the Jehol Biota, are mainly composed of volcanic-sedimentary rocks in a normal sequence. It is recommended that the Daohugou Biota and the related stratigraphy should be correlated with the Tiaojishan Formation (Lanqi Formation in West Liaoning) or its synchronous rocks. It is suggested that the Daohugou Biota and the Jehol Biota would be neither taken into one biota nor considered as the earliest elements of the Jehol Biota. The Daohugou Biota and the related rocks and the Yixian Formation were respectively formed in different periods of volcanic-sedimentary tectonics.
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A review of vertebrate fauna from the Gyeongsang Supergroup (Cretaceous) in South Korea
Article
  • Jan 2001
  • PALAEOGEOGR PALAEOCL
The Early Cretaceous is rapidly being recognized as a crucial time in the origin and dispersal of living vertebrate groups. Cretaceous trackways in Korea are among the most abundant in the world and include the smallest sauropod tracks known, plus four avian ichnotaxa, one of which is the earliest record of a bird with webbed feet. Body fossils and egg shells are less well known, however, and have been reported mainly in Korean journals.An increase in the rate of discovery of vertebrate fossils in recent years has resulted in documentation of 38 localities from the entirely fluvio-lacustrine Gyeongsang Supergroup (Hauterivian to Cenomanian) in Korea. Specimens include fish, turtle, crocodilian, pterosaur, and dinosaur bones, and dinosaur eggs in nests, as well as dinosaur, bird, and pterosaur footprints. Scattered bones have been collected in road cuts, quarries, stream beds, and coastal exposures, but localities have yet to be systematically explored and excavated. Nevertheless, very recent finds of articulated fish skeletons and dinosaur egg nests indicate that further exploration may be expected to yield better preserved, more fully associated specimens of these and additional taxa. This would provide further important data for our understanding of this pivotal period in vertebrate evolution.
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The Jehol Biota: the Emergence of Feathered Dinosaurs, Beaked Birds and Flowering Plants
  • Jan 2003
  • M M Chang
  • P J Chen
  • Y Q Wang
  • Y Wang
Chang M.M., Chen P. J., Wang Y. Q. and Wang Y. 2003. The Jehol Biota: the Emergence of Feathered Dinosaurs, Beaked Birds and Flowering Plants. Shanghai Scientific and Technical Publishers, China.
Section 5 Mesozoic Era
  • Jan 1996
  • 155-188
  • I S Pak
  • Y N Kim
Pak, I. S. and Kim, Y.N. 1996. Section 5 Mesozoic Era. In: Paek, R.J. (ed.): Geology of Korea. Publishing House, Pyongyang, pp. 155-188.
Lower Cretaceous vertebrate fauna from the Sinuiju Basin, North Korea as evidence of geographic extension of the Jehol Biota into the Korean Peninsula
  • Jan 2007
  • J Vertebr Paleontol
  • 106
  • Q Li
  • K.-Q Gao
Li, Q. and Gao, K.-Q. 2007. Lower Cretaceous vertebrate fauna from the Sinuiju Basin, North Korea as evidence of geographic extension of the Jehol Biota into the Korean Peninsula. Journal of Vertebrate Paleontology 27, supplement to number (3): 106A.
Handbuch der Paläoherpetologie, Teil 19. Pterosauria
  • Jan 1978
  • P Wellnhofer
Wellnhofer, P. 1978. Handbuch der Paläoherpetologie, Teil 19. Pterosauria, Stuttgart: Gustav Fischer Verlag.