Reassessment of the earliest documented stegosaurian fossils from Asia

Reassessment of the earliest documented stegosaurian fossils from Asia

Accepted Manuscript Reassessment of the earliest documented stegosaurian fossils from Asia Niclas H. Borinder, Stephen F. Poropat, Benjamin P. Kear PI...

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Accepted Manuscript Reassessment of the earliest documented stegosaurian fossils from Asia Niclas H. Borinder, Stephen F. Poropat, Benjamin P. Kear PII:

S0195-6671(16)30162-8

DOI:

10.1016/j.cretres.2016.08.004

Reference:

YCRES 3434

To appear in:

Cretaceous Research

Received Date: 10 May 2016 Revised Date:

1 August 2016

Accepted Date: 5 August 2016

Please cite this article as: Borinder, N.H., Poropat, S.F., Kear, B.P., Reassessment of the earliest documented stegosaurian fossils from Asia, Cretaceous Research (2016), doi: 10.1016/ j.cretres.2016.08.004. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.

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Reassessment of the earliest documented stegosaurian fossils from Asia

Museum of Evolution, Uppsala University, Norbyvägen 16, SE-752 36 Uppsala, Sweden

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Australian Age of Dinosaurs Natural History Museum, The Jump-Up, Winton, Queensland 4735, Australia

Monash University, Wellington Road, Clayton, Victoria 3800, Australia

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Niclas H. Borindera, *, Stephen F. Poropatb, c, Benjamin P. Keara

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*Corresponding author.

E-mail addresses: [email protected] (N. H. Borinder), [email protected] (Stephen F. Poropat), [email protected] (Benjamin P. Kear)

Abstract

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In 1929, the famous Swedish palaeontologist Carl Wiman documented the first unequivocal stegosaurian dinosaur fossils from Asia. His material comprised an isolated dermal spine, together with a dorsal vertebra that was briefly described but never figured. Since then these remains have languished in obscurity, being noted in some stegosaur review articles but often ignored altogether. However, recent auditing of the Museum of

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Evolution palaeontological collection at Uppsala University in Sweden has led to the rediscovery of Wiman’s original specimens, as well as two additional previously unrecognised stegosaurian dorsal vertebrae. All of these

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bones derive from the Lower Cretaceous (Berriasian–Valanginian) Mengyin Formation of Shandong Province in eastern China, and are morphologically compatible with the stratigraphically proximal stegosaurian taxon Wuerhosaurus from the Valanginian–Albian Tugulu Group in the Xinjiang Uyghur Autonomous Region of Western China. Wiman’s seminal stegosaurian fossils thus expand current palaeobiogeographical distributions, and contribute to the otherwise enigmatic record of Early Cretaceous stegosaurian occurrences globally.

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ACCEPTED MANUSCRIPT 1. Introduction Skeletal remnants of stegosaurian dinosaurs are known from every continent except Australia and Antarctica (Maidment, 2010; Pereda-Suberbiola et al., 2013). Well-preserved specimens have been known from Europe (Nopcsa, 1911a, b; Owen, 1875) North America

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(Gilmore, 1914; Marsh, 1880, 1881, 1887, 1891) and Africa (Hennig, 1915, 1925) for more than a century, with the comprehensive osteological descriptions of Stegosaurus by Gilmore (1914), and of Kentrosaurus by Hennig (1925) forming the initial basis for an understanding

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of the Stegosauria. In contrast, the earliest reports of stegosaurians from Asia were made

during the late 1920s to 1950s, and all constituted fragmentary isolated specimens (Wiman,

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1929; Young, 1935, 1944, 1958). After the 1970s, the record of Asian stegosaurian fossils improved dramatically (Dong et al., 1977; Jiang, 2006; Maidment & Wei, 2006; Zhou, 1983), and indeed, the continent now boasts more stegosaurian taxa than any other (Dong, 1990; Galton, 2012; Galton & Upchurch, 2004; Maidment, 2010; Maidment et al., 2008).

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Most Asian stegosaurian remains derive from sediments of Jurassic age, although, rare occurrences have also been documented from Lower Cretaceous deposits (Dong, 1973, 1993). The first of these discoveries was made during the Sino–Swedish expeditions of 1916-1927,

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which recovered and shipped numerous crates of dinosaur fossils from China back to Uppsala in Sweden. These now reside permanently within the Axel Lagrelius Collection of Chinese

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fossil vertebrates (Ebbestad, 2016; Mateer & Lucas, 1985) housed in the Museum of Evolution at Uppsala University (institutional acronym PMU). Much of this material derives from the Berriasian−Valanginian (Xu & Li, 2015) Mengyin Formation of Shandong Province in Eastern China (Fig. 1), which has also produced the famous titanosauriform sauropod Euhelopus zdanskyi (Mateer & McIntosh, 1985; Poropat, 2013; Poropat & Kear, 2013a; Wilson & Upchurch, 2009; Wiman, 1929). Other specimens in the collection include indeterminate titanosauriforms from the Mengyin Formation (Poropat, 2013; Upchurch &

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ACCEPTED MANUSCRIPT Mannion, 2009; Whitlock et al., 2011), as well as coelurosaurian theropods (tyrannosauroids and ornithomimosaurs: Poropat & Kear, 2013b), the ankylosaurid Pinacosaurus (Buffetaut, 1995), and the hadrosauroid ornithopod Tanius sinensis from the Upper Cretaceous Wangshi

Province (Borinder, 2015; Wiman, 1929; Xing et al. 2014).

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Group (mainly the Campanian-Maastrichtian Jiangjunding Formation), also of Shandong

Wiman’s (1929) stegosaurian specimens have attracted little recent attention (although see Pereda Suberbiola et al. [2003]), and are often overlooked in major reviews of the clade (e.g.,

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Averianov et al., 2007; Maidment, 2010; Maidment et al., 2008); nevertheless they constitute a rare and important example of unequivocal Early Cretaceous stegosaurians (Dong, 1990).

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The only earlier report of a stegosaur from Asia, that of Lametasaurus indicus from the Maastrichtian Lameta Formation of India (Matley, 1924), was later shown to be erroneous; most workers now accept that the specimen constitutes theropod remains (Chakravarti, 1935) associated with titanosaur osteoderms (D'Emic et al., 2009), although Maidment (2010),

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following Chatterjee and Rudra (1996), accepted that at least some specimens pertained to ankylosaurs. Our paper therefore reappraises Wiman’s (1929) original material as the earliest documented stegosaurian fossils from Asia. The currently registered specimens include an

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isolated dermal spine (PMU 24723) which was illustrated and described by Wiman (1929) and a dorsal vertebra (PMU 24724) that was perfunctorily described but not illustrated by

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Wiman (1929). Systematic auditing and reorganisation of the palaeontological collections at the Museum of Evolution has also uncovered two previously unrecognised stegosaurian dorsal vertebrae (PMU 20404a, PMU 20404b) by one of the authors (NHB). These bones are all assessed herein, and their palaeobiogeographical implications discussed. 2. Collection history and source rock unit According to collection archives, the PMU Chinese stegosaurian remains were recovered in 1922 by H. C. T’an, a geologist with the National Geological Survey of China, and Johan

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ACCEPTED MANUSCRIPT Gunnar Andersson, the expedition leader for the Sino–Swedish field exploration programme,

who later became famous as “Kina-Gunnar” (Mateer & Lucas, 1985). The specimen labels list “Ning-Chia-Kou, Meng-Yin Hsien, Shantung” as the source locality. Today this corresponds to the village of Ningjiagou in Mengyin County, western central Shandong Province (Fig. 1A,

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B). Unfortunately no field notes and only a few site photos have survived (see Fig. 1C);

however, the dorsal vertebrae PMU 20404a and PMU 20404b were stored with a common label, and their morphology (e.g. size, ontogeny) and preservation (e.g., adhering grey–grey-

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green sandstone and red oxidized mineral staining) suggests potential anatomical association. Wiman (1929, 1930) and Stensiö (1935) otherwise attributed all of the PMU Ningjiagou

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dinosaur remains to the lower and middle horizons of the Mengyin Formation. This was based on matrix compatibility, which we confirmed here via first-hand inspection of sediments encasing indisputable Mengyin Formation fossils such as the referred pelvic elements of

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Euhelopus zdanskyi (PMU 24706).

3. The Mengyin Formation fossil biota

The Mengyin Formation has yielded numerous vertebrate fossils. Most notable are the two

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partial skeletons and referred elements of Euhelopus zdanskyi (Mateer & McIntosh, 1985; Poropat, 2013; Poropat & Kear, 2013a; Wilson & Upchurch, 2009; Wiman, 1929; Young,

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1935), which were found within a few kilometers of Ningjiagou. Euhelopus-like teeth have been reported from the Aptian Yixian (Barrett & Wang, 2007) and Shahai formations (Amiot et al., 2010). Other reported dinosaur remains are fragmentary. For example, Young (1935) described the right scapula of a sauropod (recovered ~250 m northwest of “Hsichüfu”) and the dermal spike of a stegosaurian, excavated ~3 km northwest of “Hsichüfu” that closely resembled PMU 24723 (although considerably larger).

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ACCEPTED MANUSCRIPT Young (1935) also reported an isolated right coracoid, found near Ningjiagou, which he assigned to Sauropoda (reassigned to Theropoda by Wilson & Upchurch [2009]), and an ?ulna, found ~2.5 km west of Hsichüfu, which he assigned to Theropoda (reinterpreted as a pterosaur wing phalanx by Wilson & Upchurch [2009]).

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The cryptodiran turtles Sinemys lens (Brinkman & Peng, 1993; Wiman, 1930) and

Sinochelys applanata (including Scutemys tecta: Čkhikvadze, 1985; Wiman, 1930), together with the amiiform fish Sinamia zdanskyi (Liu & Su, 1983; Stensiö, 1935) and

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osteoglossomorph Lycoptera (Stensiö, 1935; Wiman, 1929; Young, 1935; Zhang, 2012) are represented by abundant material. The crocodylomorph Shartegosuchus chuhsienensis might

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also derive from the Mengyin Formation (Young, 1961). Wu et al. (1994) cited the presence of Sinemys and Shartegosuchus as correlates for an Early Cretaceous age because of their occurrence elsewhere within the Barremian (or younger) Luohandong Formation of Inner Mongolia. Finally, sinamiids including Sinamia (Cavin et al., 2007; Peng et al., 2015;

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Yabumoto, 2014; Zhang, 2012), and the sinemydid group incorporating Sinemys (Brinkman & Peng, 1993; Tong & Brinkman, 2013) are widespread throughout Lower Cretaceous

Korea and Japan.

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(Barremian–Albian) units in northeastern–southeastern Asia from China, to Thailand, South

Grabau (1923) described various freshwater molluscs from the Mengyin Formation

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(likewise briefly mentioned by Stensiö, 1935; Wiman, 1929, 1930), such as the unionid bivalve Mycetopus mengyinensis (generically reassigned to Mengyinaia and Margaritifera, including a second species M. tugrigensis: Chen, 1984; Gu et al., 1976 in Sha et al., 2006; Ma, 1996 in Pan & Sha, 2009) and heterobranch gastropods Valvata suturalis (= Amplovalvata suturalis: Pan, 1983) and Bithynia mengyinense. Ma (1994) assigned the bivalve assemblage a Berriasian–Barremian age range. Chen (1982) additionally reported the conchostracan Eosestheria, which occurs throughout the Lower Cretaceous of China (Li et al.,

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ACCEPTED MANUSCRIPT 2007). The ubiquitous non-marine ostracod Cypridea has also been recorded from the Mengyin Formation (Zhang et al., 2003), along with indeterminate plant fragments (Stensiö,

4. Systematic Palaeontology DINOSAURIA Owen 1842 ORNITHISCHIA Seeley 1887

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THYREOPHORA Nopcsa 1915 sensu Norman 1984

4.1. Referred material

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STEGOSAURIA Marsh 1877

Stegosauria gen. et sp. indet.

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1935; Wiman, 1929).

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PMU 24724 (formerly PMU R275), PMU 20404a and PMU 20404b: three dorsal vertebrae. PMU 24723 (formerly PMU R276): an isolated dermal spine.

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4.2. Locality and horizon

PMU 20404a, PMU 20404b, PMU 24724 and PMU 24723 were all recovered in the

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vicinity of Ningjiagou village (“Ning-Chia-Kou, Meng-Yin Hsien” and “near Ho Tung, Nan Ling”: Wiman, 1929), Mengyin County in western central Shandong Province (Fig. 1A–B). The source horizon was almost certainly within the Mengyin Formation (Chen et al., 1980; see Xu & Li, 2015, p. 285, fig. 1 for a detailed geological map). The age of this unit has been contested (see Wilson & Upchurch, 2009) but might be Berriasian–Barremian (Ma, 1994) or possibly Aptian (Barrett & Wang, 2007) based on faunal correlations. Recently, detrital zircon

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ACCEPTED MANUSCRIPT and U-Pb zircon dating constrained this range to ca. 145–136 Ma, corresponding to the Berriasian–Valanginian (basal Early Cretaceous; Xu & Li, 2015).

5. Description and comparisons

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5.1. Vertebrae

PMU 20404a, PMU 20404b and PMU 24724 incorporate intact centra with the neural arches broken off below the zygapophyses and transverse process bases (Figs. 2A–J; 3A–D). These

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natural breaks reveal dense internal bone structure, which contrasts with the usually highly pneumatized bone of saurischian dinosaurs (Britt, 1993, 1997; Wedel, 2006). The centra are

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amphicoelous, with oval articular faces that are transversely narrower than dorsoventrally high (Table 1). The facet surfaces on PMU 20404a and PMU 20404b are alternately concave versus flattened on the opposing face (Figs. 2A, C, F, H); this resembles the anterior dorsal vertebrae of stegosaurians (e.g. Alcovasaurus longispinus: Gilmore, 1914; Galton &

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Carpenter, 2016). In contrast, the centrum of PMU 24724 is proportionately larger and anteroposteriorly elongate relative to its maximum dorsoventral height (see Table 1). This might place PMU 24724 within the mid-dorsal series based on comparisons with Stegosaurus

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stenops; in this taxon the centrum length progressively increases along the mid-dorsal region, before decreasing again towards the tail (Maidment et al., 2015). A similar trend has been

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reported in Kentrosaurus aethiopicus (Hennig, 1925; Mallison, 2010), Loricatosaurus priscus (Galton, 1985, 1990), Chialingosaurus kuani (Carpenter et al. 2001), and Huayangosaurus taibaii (Maidment et al., 2006), but not in Dacentrurus armatus (Owen, 1875, pl. 13; Galton, 1985; Maidment et al., 2008). The lateral surfaces of the centra are deeply concave, imparting a constricted “hourglassshape” in ventral view (Figs. 2E, J; 3D); this is common amongst stegosaurians (Galton &

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ACCEPTED MANUSCRIPT Upchurch, 2004). The articular facet rims are thus projected, and the dorsal edges of the centra indent to accommodate the floor of the neural canal. Only the base of the neural arch is preserved in PMU 20404a, PMU 20404b and PMU

24724, but the neurocentral sutures are all fully fused (although still traceable on PMU 24724:

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Fig. 3B) implying osteological maturity. The neural arches are transversely compressed

(especially in PMU 24724), and were apparently tall, as is typical of stegosaurians (Galton & Upchurch, 2004) including Alcovasaurus and Stegosaurus stenops (Gilmore, 1914; Maidment

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et al., 2015), Kentrosaurus (Hennig, 1925), Loricatosaurus (Galton, 1985, 1990),

Chialingosaurus (Carpenter et al., 2001), and Huayangosaurus (Maidment et al., 2006).

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However, PMU 20404a, PMU 20404b and PMU 24724 differ from Huayangosaurus (Maidment et al., 2006), Hesperosaurus mjosi, (Carpenter et al., 2001; Maidment et al., 2015) and Gigantspinosaurus sichuanensis (Peng et al., 2005; Maidment et al., 2015) in that their prezygapophyses are elevated well above the neural canal.

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In PMU 24724 the neural canal is transversely narrow and ‘teardrop-shaped’ to elliptical (although this was likely modified by diagenetic distortion: Fig. 3A, C). On the other hand, the neural canals of PMU 20404a and PMU 20404b are almost circular in outline (Fig. 2A, C,

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F, H). The vertical surface immediately above the neural canal is indented to form a shelf. Blunt protuberances also project from the lateral margins of the neural canal opening above

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the pedicles; these are most evident on PMU 24724 (Fig. 3A–C). The neural arches of PMU 20404a, PMU 20404b and PMU 24724 all bear deep fossae that incise the vertical surface above the neural canal on at least one side (possibly the anterior) in PMU 20404a (Fig. 2C) and PMU 24724 (Fig. 3C), and on both opposing surfaces of the neural arch in PMU 20404b (Fig. 2F, H). Maidment et al. (2008, p. 379) listed similar “[deep excavations] dorsal to the neural canal in anterior view” (Fig. 4H–N) as diagnostic for Wuerhosaurus homheni (reassigned to the genus Stegosaurus by Maidment et al., 2008).

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ACCEPTED MANUSCRIPT Maidment et al. (2008, p. 380, fig. 8A) also labeled a corresponding “pathological excavation” on the anterior face of a dorsal vertebra recovered with the W. homheni holotype specimen (Institute of Vertebrate Paleontology and Paleoanthropology [IVPP], Beijing IVPP V4006: Dong, 1973). Based on photographs (images supplied by pers. comm. S. Maidment,

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2016: see Fig. 4A–G), comparable fossae are present on the dorsal vertebrae of Wuerhosaurus ordosensis (e.g., IVPP V6879: Dong, 1993), and offer a means for orienting the left and right sides of PMU 20404a, PMU 20404b and PMU 24724. The opposite dorsalmost face of the

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neural arches on PMU 20404a and PMU 24724 are otherwise flat, with a prominent ridge extending vertically along the midline (this compares well with the ridges extending vertically

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from the zygapophyses in S. stenops: see Maidment et al., 2015). Weak parallel ridges are also present in PMU 24724 (Fig. 3A), and a comparable structure traverses obliquely across the fossa on PMU 20404a (Fig. 2A).

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5.2. Dermal spine

Wiman (1929) identified PMU 24723 as a stegosaurian dermal spine, probably derived from the right side of the body. The element is incomplete, missing the distal third or half (Fig. 5A–

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D). Its base is rhombic in proximal outline (Fig. 5E), and obliquely offset such that the proximomedial margin expands asymmetrically (Fig. 5D). The spine itself is thus acutely

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inclined away from the vertical axis, which probably led Wiman (1929) to orient it towards the right-hand side. The proximal edges are rugose with a crenellated edge. The distal extremity of the spine is straight, rhombic and transversely narrowed in cross-section, with prominent tapered edges along its anterior and posterior sides (Fig. 5B, D). Although PMU 24723 superficially resembles the caudal spines of Stegosaurus ungulatus (Gilmore, 1914), Hesperosaurus mjosi (Carpenter et al., 2001) and Stegosaurus stenops (Maidment et al., 2015), it is much smaller (see Table 2), and lacks the massively expanded proximal base (see

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ACCEPTED MANUSCRIPT Kentrosaurus aethiopicus: Galton, 1982). Gilmore (1914) noted that the caudal spines of

Stegosaurus were highly variable in size and shape. Therefore we consider PMU 24723 to be derived either from the tail region or caudal end of the trunk, and we agree with Wiman

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(1929) in that it is from the right side of the body.

6. Discussion

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Attribution of the PMU Mengyin Formation dorsal vertebrae (PMU 20404a, PMU 20404b, PMU 24724) and dermal spine (PMU 24723) to Stegosauria is based on overall

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morphological compatibility. In particular, the transversely compressed and dorsoventrally high vertebral proportions, incorporating zygapophyses that are elevated high above the neural canal, serves to differentiate the PMU specimens from ankylosaurians. These usually possess dorsal centra that are transversely broader than long (Vickaryous et al., 2004), and

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pre/postzygapophyses that are situated immediately proximal to the neural canal roof – e.g., Hungarosaurus tormai (Ősi, 2005), Peloroplites cedrimontanus (Carpenter et al., 2008), Europelta carbonensis (Kirkland et al., 2013), Euoplocephalus tutus (Arbour & Currie, 2013),

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and Polacanthus foxii (Blows & Honeysett, 2014). In addition, ankylosaurian dermal spines tend to be more conical (e.g., P. foxii and species of Struthiosaurus: Blows, 1987; Ősi, 2015),

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or flattened and triangular with a broad base and occasional bifurcation (e.g., Sauropelta edwardsorum, Edmontonia rugosidens: Carpenter, 1984, 1990). Moreover, ankylosaurian pectoral spines can exhibit conspiucuous grooves or fluting (e.g., Gastonia burgei, Hylaeosaurus armatus: Kirkland, 1998; Sachs & Hornung, 2013). Finally, the coeval Chinese ankylosaurians including Dongyangopelta yangyanensis, Taohelong jinchengensis and Zhejiangosaurus lishuiensis are not known to possess dermal spines (Lü et al., 2007; Chen et al., 2013; Yang et al., 2013); however, each of these taxa is represented by incomplete and

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ACCEPTED MANUSCRIPT scattered postcranial remains pertaining to the posterior half of the body – the least likely

region for spikes based on the anatomy of other ankylosaurs. The morphology of PMU 24723 is compatible with that of stegosaur spikes, prompting us to assign the spike to Stegosauria rather than Thyreophora indet. (e.g., Maidment et al., 2008).

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The most compelling argument, however, for referral of this material to Stegosauria is the presence of deep vertical excavations on the dorsal neural arches of PMU 20404a, PMU

20404b and PMU 24724. This feature directly complies with the diagnosis of Wuerhosaurus

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homheni (Fig. 4H–N) from the ?Valanginian−Albian Tugulu Group (Lianmuging Formation) of the Xinjiang Uyghur Autonomous Region in Western China (Maidment et al., 2008). A

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second species, W. ordosensis (Fig. 4A–G; considered a nomen dubium by Maidment et al., 2008) from the Aptian-Albian Ejinhoro Formation of Inner Mongolia (Nei Mongol) (Galton & Upchurch, 2004) also manifests this trait. Our identification of comparable stegosaurian remains from the Mengyin Formation, together with stegosaurian-like footprints identified as

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cf. Deltapodus from the upper Tuchengzi Formation (= Berriasian: Xu & Li, 2015) near Beijing in eastern China (Zhang et al., 2012), thus collectively suggests that this group formed a widespread component of Early Cretaceous dinosaurian faunas in Asia. Indeed, Deltapodus

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curriei tracks have been reported along with possible limb bones of Wuerhosaurus from the Tugulu Group (Xing et al., 2013), and the Late Jurassic–Early Cretaceous stegosaurian taxon

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Monkonosaurus lawulacus is also known from the Loe-ein Formation of Tibet (Dong, 1990; Maidment et al., 2006). The highly controversial Dravidosaurus blanfordi from the Coniacian Trichinopoly Group of India (Yadagiri & Ayyasami, 1979) has been reinterpreted as a probable plesiosaurian (see Maidment et al., 2008 and references therein), but Deltapoduslike thyreophoran footprints are nonetheless recognized from the Maastrichtian Infratrappean Limestone of Gujarat of India (Mateus et al., 2011; Mohabey, 1986). In conjunction with occurrences from Europe (Alcalá et al., 2014; Billon-Bruyat et al., 2010; Cobos et al., 2010;

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ACCEPTED MANUSCRIPT Company et al., 2010; Maidment, 2010; Pascual et al. 2012; Pascual-Arribas & HernándezMedrano 2015; Weishampel et al., 2004), Africa (Maidment et al., 2008), South America

(Apesteguía & Gallina, 2011; Pereda-Suberbiola et al., 2013), and possibly Australia (Milàn & Chiappe 2009), the PMU Mengyin Formation specimens contribute to the growing picture

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of Early Cretaceous stegosaurians as a globally distributed clade (Fig. 6). Their apparent rarity is therefore likely an effect of sampling, but as shown here, historical museum collections might provide an important source of undiscovered remains in the future.

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7. Conclusions

•The earliest documented Asian stegosaurian fossils were recovered during the Sino-Swedish

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expeditions of 1916−1927, and derived from the Lower Cretaceous (Berriasian–Valanginian) Mengyin Formation of Shandong Province in eastern China. The specimens include a dorsal spine and three dorsal vertebrae that are morphologically consistent with the approximately coeval (?Valanginian−Albian) Chinese stegosaurian taxon Wuerhosaurus.

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•The demonstrable occurrence of stegosaurians in the Mengyin Formation dinosaurian assemblage indicates a widespread palaeobiogeographical distribution throughout the Lower Cretaceous of China. This contrasts with other regional faunas, which infer extinction of

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stegosaurians at the Jurassic–Cretaceous boundary (e.g. North America: Galton & Upchurch, 2004; Maidment et al., 2008) but might be reflective of stratigraphical bias versus the much

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richer Lower Cretaceous records of Asia and Europe (Tennant et al., 2016).

Acknowledgements

Thanks to Jan Ove Ebbestad (Museum of Evolution) for specimen preparation and collections management. Susannah Maidment (Imperial College) generously contributed both photographs and information, and together with an anonymous reviewer provided helpful comments which improved the manuscript.

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ACCEPTED MANUSCRIPT PMU 24724

PMU 24040a

PMU 24040b

Centrum length

82.41

54.64

56.61

Centrum articular facet dorsoventral height/width

56.34/55.07

47.78/52.74

49.62/53.76

Centrum articular facet dorsoventral height/width

58.10/59.35

44.71/53.47

48.24/51.39

Neural canal diameter proximal/distal

15.98/14.61

22.08/18

21.66/18.93

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Specimen

Table 1. Measurements (mm) of PMU stegosaurian dorsal vertebrae from the Mengyin Formation, Ningjiagou,

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Shandong, China.

PMU 24723

Proximodistal length

119.81

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Maximum length/width at base

64.55/39.37

Table 2. Measurements (mm) of PMU stegosaurian dermal spine from the Mengyin Formation, Ningjiagou,

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Shandong, China.

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Figure 1. A, Map of China with (B) enlargement of Shandong Province indicating Ningjiagou village (Mengyin County) and other adjacent Cretaceous vertebrate fossil localities. C, field photograph by Johan Gunnar Andersson likely showing the Mengyin excavation area (Andersson, 1923).

lateral; C/H, articular; D/I, right lateral; E/J ventral views. Scale bar = 50 mm.

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Figure 2. Stegosaurian dorsal vertebrae PMU 24040a (A–D) and PMU 24040b (F–J). A/F, articular; B/G, left

Figure 3. Stegosaurian dorsal vertebra PMU 24724. A, articular; B, right lateral; C, articular; D, ventral views.

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Scale bar = 50 mm.

Figure 4. Figure 4. Wuerhosaurus ordosensis dorsal vertebra IVPP V6879 (A–G) and Wuerhosaurus homheni

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dorsal vertebra IVPP V4006 (H–N). A–B/H–I, anterior; C–D/J–K, left lateral; E–F/L–M, posterior; and G/N, right lateral views. (All photographs courtesy S. Maidment [pers. comm. 2016]). Scale bar = 100 mm.

Figure 5. Stegosaurian dermal spine PMU 24723. A, lateral; B, anterior; C, medial; D, posterior; E,

50 mm.

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proximalviews (based on orientation towards the right-hand side of the body: sensu Wiman, 1929). Scale bar =

Figure 6. Early Cretaceous palaeogeographical map showing distribution of stegosaurian skeletal fossils (body

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outline) and footprints (track outline). Modified from Pereda-Suberbiola et al. (2013). 1. China: Mengyin Formation, Shandong Province (this study). 2. China: Tugulu Group, Xinjiang Uyghur Autonomous Region

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(Galton & Upchurch, 2004). 3. China: Tugulu Group, Xinjiang Uyghur Autonomous Region (Xing et al., 2013). 4. China: Ejinhoro Formation, Inner Mongolia Autonomous Region (Galton & Upchurch, 2004). 5. Tibet: Loeein Formation, Markam (Dong, 1990). 6. China: Tuchengzi Formation, Beijing (Zhang et al., 2012). 7. France: unnamed unit, Charente (Billon-Bruyat et al., 2010). 8. England: Potton Sands, Bedfordshire; Wealden Group, Sussex, (Maidment, 2010; Weishampel et al., 2004). 9. Spain: Villar del Arzobispo Formation, various localities in Teruel and Valencia; El Castellar Formation, Teruel; Piedrahita de Muñó Formation, Burgos; Camarillas and Artoles formations, Teruel (see Company et al., 2010 for summary). 10. Spain: Villar del Arzobispo Formation, Teruel; Oncala Group, Soria; Magaña Formation, Soria (Cobos et al., 2010; Pascual et al., 2012; Pascual-Arribas & Hernández-Medrano 2015). 11. Argentina: La Amarga Formation, Neuquén (Pereda-Suberbiola et al., 2013).

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Formation, Algoas Basin (Maidment et al., 2008).14. Australia: Broome Sandstone, Western Australia (Milàn &

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