Zootaxa 3914 (4): 421–440
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ArticleISSN 1175-5326 (print edition)ZOOTAXAISSN 1175-5334 (online edition)

http://dx.doi.org/10.11646/zootaxa.3914.4.3
http://zoobank.org/urn:lsid:zoobank.org:pub:155185CB-A231-47E6-ACC9-39E11FCDDFA5

New basal hadrosauroid (Dinosauria: Ornithopoda) from the Lower Cretaceous
Kitadani Formation, Fukui, central Japan
MASATERU SHIBATA1,2 & YOICHI AZUMA1,2
1

Dinosaur Research Institute, Fukui Prefectural University, 4-1-1 Kenjojima, Matsuoka, Eiheiji-Cho, Fukui, 910-1195 Japan.
E-mail: sdino@fpu.ac.jp, y-azuma@fpu.ac.jp
2
Fukui Prefectural Dinosaur Museum, 51-11, Terao, Muroko, Katsuyama-Shi, Fukui 911-8601, Japan

Abstract
A new basal hadrosauroid, Koshisaurus katsuyama, from the Lower Cretaceous Kitadani Formation in Fukui, central Japan, is reported. The new taxon is distinguished by an autapomorphy and a unique combination of maxillary, vertebral,pubic and femoral characters. A phylogenetic analysis indicates that Koshisaurus is positioned as a basal member of Hadrosauroidea and is more derived than the contemporaneous Fukuisaurus, which is a non-hadrosauroid hadrosauriform.
The presence of the antorbital fossa on the maxilla and at least three subsidiary ridges on the labial side of maxillary toothcrown implies that Koshisaurus was among the most basal hadrosauroids. This discovery indicates a higher diversity ofhadrosauroids along the eastern margin of the Asian continent in the Early Cretaceous.
Key words: Early Cretaceous, Hadrosauroidea, Kitadani Formation, Tetori group, Japan

Introduction
Recent discoveries and descriptions of styracosternan iguanodonts indicate the great diversity of this groupthroughout the Early Cretaceous (You & Li 2009; McDonald et al. 2010a, b; Wu et al. 2010; You et al. 2011;
McDonald et al. 2012; Wu & Godefroit 2012). In Asia, the fossil record of this group is extensive. However,discoveries in Asia are largely restricted to dinosaur-rich formations in China and Mongolia. Until recently,
Fukuisaurus tetoriensis from the Lower Cretaceous (Barremian–lower Aptian) Kitadani Formation of Japan was
the only named Asian styracosternan recognized from outside of this region (Kobayashi & Azuma 2003). The
recent reports of two species of styracosternans in Thailand extended their diversity and distribution into Southeast
Asia in the late Early Cretaceous (Buffetaut & Suteethorn 2011; Shibata et al. 2011). In Japan, styracosernan
iguanodont discoveries in addition to Fukuisaurus include a maxillary tooth from the Tatsukawa Formation(Hauterivian) in Tokushima, several well-preserved teeth from the Kuwajima Formation (Barremian) in Ishikawa(Morozumi et al. 1995; Manabe & Barrett 2000), and shed teeth from the “Lower Formation” (lower Albian) of the
Sasayama Group in Hyogo (Saegusa & Ikeda 2014).
The Kitadani Formation, the uppermost unit of the Tetori Group, is the most productive dinosaur-bearing unit
in Japan, having yielded Fukuiraptor kitadaniensis, Fukuisaurus tetoriensis and Fukuititan nipponensis (Azuma &Currie 2000; Kobayashi & Azuma 2003; Azuma & Shibata 2010). Additionally, ornithopod dentaries, which do not
pertain to Fukuisaurus, have also been reported (Shibata & Azuma 2011). All of these dinosaurian specimens were
found in one locality, the Kitadani Quarry in Katsuyama City (Fig. 1). At least four bonebeds are recognized at thislocality (Fig. 2A); the definition of “bonebed” in this study follows Behrensmeyer (2007). The lowermost one, theBonebed I (BB I), has yielded Fukuiraptor, Fukuisaurus, other dinosaur bones and teeth, and other vertebrates
including crocodilians, turtles and fishes (Shibata & Goto 2008). Most of the BB I vertebrate assemblage consistsof disarticulated or associated skeletons of dinosaurs and other vertebrates. Other horizons (BB II and III), which
are approximately 10 m above BB I, yielded Fukuititan elements and a well-preserved small theropod skeleton(Shibata & Goto 2008), respectively. The iguanodontian bones described in this paper came from the fourth bone

Accepted by S. Brusatte: 16 Dec. 2014; published: 29 Jan. 2015421

In all consensus trees the two Fukui taxa are not particularly closely related. In the Adams consensus tree, forexample, Koshisaurus is positioned at the base of Hadrosauroidea. In contrast, the other Fukui iguanodontian,
Fukuisaurus, falls out as a basal hadrosauriform and closely related to Proa, Iguanodon and Ouranosaurus.
Consequently, our analysis corroborates that Koshisaurus and Fukuisaurus are phylogenetically distinct;
Koshisaurus is more derived than Fukuisaurus. Coexistence of at least two taxa of styracosternans occurred
repeatedly during the Early Cretaceous. Examples include Barilium and Hypselospinus in the Wadhurst Clay
Formation of England, Cedrorestes, Iguanocolossus and Hippodraco in the Cedar Mountain Formation of Utah,
Iguanodon and Mantellisaurus in the Wessex Formation of England and Sainte-Barbe Clays Formation ofBelgium, Bolong and Jinzhousaurus in the Yixian Formation of China, and Lurdusaurus and Ouranosaurus in the
El Rhaz Formation of Niger (Taquet 1976; Norman 1980, 1986; Taquet & Russell 1999; Wang & Xu 2001;You and
Ji et al. 2003; McDonald et al. 2010b; Wu & Godefroit 2012).
Koshisaurus was one of the most primitive forms of Hadrosauroidea because it possesses an antorbital fossa
and three subsidiary ridges on the maxillary teeth, similar to Equijubus (You et al. 2003). The subsidiary ridges ofthe maxillary teeth are also present in Xuwulong, Jinzhousaurus and Altirhinus among basal hadrosauroids, butthese taxa lack the antorbital fossa on the maxilla (Norman 1998; You et al. 2003; Barrett et al. 2009).

Conclusions
A new hadrosauroid, Koshisaurus katsuyama from the Early Cretaceous Kitadani Formation, is a distinct new
genus and species and distinguishable from the contemporaneous hadrosauriform Fukuisaurus tetoriensis. Due toits possession of an antorbital fossa and at least three subsidiary ridges on the crown of the maxillary teeth,
Koshisarus is found to be one of the most primitive hadrosauroids in our phylogenetic analysis. The occurrence ofnumerous basal hadrosauroids in Asia suggests a diversification of this group in Asia during the Early Cretaceous
period.

Acknowledgments
We are very thankful to all the people who contributed to the 3rd Dinosaur Excavation Project of Fukui Prefecture
2007–2010 and to the preparators at FPDM for their patient and skillful work. We are grateful to Drs. David
Varricchio (Montana State University, Bozeman), Joshua Bonde (University of Nevada, Las Vegas), Kyo Tanoue(Fukuoka University, Japan), and Mr. Wenjie Zheng (Zhejiang Museum of Natural History, China) for their advice
and editing this manuscript. One of authors (MS) extends special appreciation to Dr. Jack Horner (Museum of theRockies, Montana State University, Bozeman), who encouraged the sampling of this material for histology, and to
the Paleohistology Lab manager of MOR, Ellen-Therese Lamm for teaching the techniques for histologicalsectioning. Finally, we greatly appreciate useful comments and suggestions by the editor and reviewers to improve
this manuscript. The Fukui Prefectural Government supported this research program.

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