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Cretaceous Research 45 (2013) 103e113

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Cretaceous Research

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Reassessment of coelurosaurian (Dinosauria, Theropoda) remainsfrom the Upper Cretaceous Wangshi Group of Shandong Province,China

Stephen F. Poropat a,b,*, Benjamin P. Kear a

aDepartment of Earth Sciences, Uppsala University, Uppsala 752 36, SwedenbAustralian Age of Dinosaurs Museum of Natural History, The Jump-Up, Winton, Queensland 4735, Australia

a r t i c l e i n f o

Article history:Received 18 June 2013Accepted in revised form 11 August 2013Available online

Keywords:TheropodaCoelurosauriaTyrannosauroideaOrnithomimosauriaChinaCretaceous

* Corresponding author. Department of Earth SUppsala 752 36, Sweden.

E-mail addresses: stephen.poropat@geo.uu.se,(S.F. Poropat), benjamin.kear@geo.uu.se (B.P. Kear).

0195-6671/$ e see front matter � 2013 Elsevier Ltd.http://dx.doi.org/10.1016/j.cretres.2013.08.005

a b s t r a c t

Non-avian theropods are well represented in the Cretaceous of Asia. However, the first theropod re-mains ever reported from China, from the Wangshi Group of Shandong Province, have not beenreassessed since they were described by Carl Wiman in 1929, despite the great strides that have beenmade in theropod phylogenetics in the last eight decades. The remains redescribed herein consist offour vertebrae (two cervicals, one dorsal and one caudal) from one site, and a caudal vertebra andungual phalanx from another. The vertebrae from the first site are assigned to tyrannosauroids andornithomimosaurs, as is the caudal vertebra from the second site, whereas the manual ungual alsobears resemblances to those of non-maniraptoran coelurosaurs (specifically compsognathids andornithomimosaurs). Theropods remain relatively rare in the Wangshi Group; nevertheless, the remainsredescribed herein highlight the potential for the discovery of more theropod remains in futureexcavations.

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1. Introduction

Non-avian theropod dinosaurs are well represented in theMesozoic rocks of China. The Lower Cretaceous Jehol (including theYixian and Jiufotang formations) and Xinminpu groups, and theUpper Cretaceous Wulansuhai Formation, have all preserveddiverse theropod records, as have several other formations(Weishampel et al., 2004). In contrast to these productive rockunits, and those of the Upper Cretaceous of Mongolia, the UpperCretaceous Wangshi Group of Shandong Province (Fig. 1) has yiel-ded relatively few theropod specimens, despite having producedremains of indeterminate cerapodans (Butler and Zhao, 2009;Dong, 1978), abundant hadrosaurs (Hu, 1973; Hu et al., 2001; Jiet al., 2011; Wiman, 1929; Young, 1958; Zhao et al., 2011; Zhaoand Li, 2008; Zhao et al., 2007), ceratopsians (Xu et al., 2010a,b),ankylosaurs (Buffetaut, 1995; Gilmore, 1933), and sauropods(Poropat, 2013; Wiman, 1929). Following Hu et al. (2001), the

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Wangshi Group comprises four formations; the theropod remainsfrom each are summarised below.

The oldest formation within the Wangshi Group, the Xingez-huang Formation, was reported by Hu et al. (2001) to containtheropod remains, specifically a metatarsal (GMC V1777) and teeth(GMC V286, V288, V1174 and V1773), which were originally likenedto Tyrannosaurus rex Osborn, 1905 (Hu, 1973) but later assigned tothe new species Tyrannosaurus zhuchengensis Hu, Cheng, Pang, andFang, 2001. Themetatarsal was identified as a right metatarsal IV byHu (1973) and Hu et al. (2001) but was reinterpreted as a probableleft metatarsal II by Hone et al. (2011), who designatedT. zhuchengensis as a nomen dubium. In the same work, Hone et al.(2011) erected the new taxon Zhuchengtyrannus magnus Hone,Wang, Sullivan, Zhao, Chen, Li, Ji, Ji, and Xu, 2011 on the basis ofan associated rightmaxilla and left dentary (ZCDMV0031) from theXingezhuang Formation near Zhucheng.

The Jiangjunding Formation, fromwhich Tanius sinensisWiman,1929 was recovered, was also said to have yielded theropod re-mains by Hu et al. (2001). Hu et al. (2001) were referring to thetheropod specimens described by Wiman (1929), collected fromtwo separate localities which also contained Tanius sinensis. Theseremains, which have not been discussed since their originaldescription by Wiman (1929), form the basis of this paper (seebelow).

Fig. 1. Map of China with an enlarged map of Shandong Province (inset) showing several towns near which Cretaceous dinosaur fossils have been found. The theropod remainsdescribed in this paper come from localities within a few kilometres of Jingangkou. Scale bar ¼ 100 km.

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The presence of ‘carnosaur’ material in the Jingangkou Forma-tion was noted by Hu et al. (2001). This material includes: Ching-kankousaurus fragilis Young, 1958, a tyrannosauroid known onlyfrom an incomplete scapula (Brusatte et al., 2013); the ‘carnosaur’material described by Young (1958) as being similar to the mucholder Szechuanosaurus campi Young, 1942; and a large theropoddorsal vertebra (centrum length 91 mm; vertebra height 310 mm)also described by Young (1958).

The youngest formation in theWangshi Group, the ChangwanpuFormation, has not yet yielded any theropod material, according toHu et al. (2001).

The theropod remains redescribed in this paper were the firstever reported from China, initially described by Carl Wiman (1929)of the Palaeontological Museum, Uppsala, Sweden (PMU). Thesefossils were collected in 1923 by C. H. T’an of the Chinese GeologicalSurvey, and Otto Zdansky, an employee of the PMU, from twoseparate localities in Shandong (Shantung) Province, China(Table 1). The theropod remains consist of four vertebrae (twocervicals, one dorsal, and one caudal) found at the same site as theholotype specimen of Tanius sinensis, and a claw and a caudalvertebra from a different site. Unfortunately, Zdansky’s field note-books were stolen on the Trans-Siberian Express as he made his

way back to Sweden (J. O. R. Ebbestad, pers. comm., March, 2013),meaning that there is now no way to determine if the remains ateach sitewere associated. Therefore, in this work, wewill treat eachspecimen as an isolated element, even when others in the collec-tion are known to have come from the same locality (Table 1).

Institutional Abbreviations. BSP, Bayerische Staatsammlung für Pal-äontologie und historische Geologie, Munich, Germany; GIN,Paleontological Center of Mongolia, Ulaan Bataar, Mongolia; GMC,Geological Museum of China; PMU, Palaeontological Museum,Uppsala, Sweden; ZCDM, Zhucheng Dinosaur Museum, China.Anatomical Abbreviations. acdl, anterior centrodiapophyseal lamina;cdf, centrodiapophyseal fossa; cpof, centropostzygapophysealfossa; cpol, centropostzygapophyseal lamina; cprf, centroprezygapophyseal fossa; dp, diapophysis; f, fossa; ha, hypantrum; hs,hyposphene; k, keel; nc, neural canal; ns, neural spine; pcdl, pos-terior centrodiapophyseal lamina; pcpl, posterior centropar-apophyseal lamina; pfr, pneumatic foramen; pfs, pneumatic fossa;pocdf, postzygapophyseal centrodiapophyseal fossa; podl, post-zygodiapophyseal lamina; posf, postspinal fossa; poz, post-zygapophysis; pp, parapophysis; prcdf, prezygapophysealcentrodiapophyseal fossa; prdl, prezygodiapophyseal lamina; prsf,

Table 1Theropod specimens from the Upper Cretaceous Wangshi Group of Shandong Province, eastern China, held in the collections of the Palaeontological Museum, Uppsala,Sweden.

Locality data Specimen number Old specimen number Description

20 Apr. 1923 H.C. T’anLai-Yang-HsienChiang Chün TingShantungSW 1 li

PMU 24711 e Cervical vertebra (partial)PMU 24712 e Cervical vertebraPMU 24713 PMU R 270 (& 219(?)) Dorsal vertebraPMU 24714 PMU R 279 & 285 Caudal vertebraPMU 24715 PMU R 278 Indeterminate fragmentPMU 24716 PMU R 280, 281, 283 & 284 Indeterminate fragmente PMU R 282 Indeterminate fragment; may attach to another specimen

Okt. 1923 ZdanskyShantungLai-Yang-HsienS 16 liChiang-Chün-TingNW 1 li

PMU 24717 PMU R 273 Manual ungual phalanxPMU 24718 PMU R 274 Vial of shattered fragments; may have been a toothPMU 24719 PMU R 277 Caudal vertebra

S.F. Poropat, B.P. Kear / Cretaceous Research 45 (2013) 103e113 105

prespinal fossa; prz, prezygapophysis; spof, spinopostzygapophy-seal fossa; spol, spinopostzygapophyseal lamina; sprl, spinoprezy-gapophyseal lamina; tpol, intrapostzygapophyseal lamina; tprl,intraprezygapophyseal lamina.

2. Descriptions

All of the vertebrae discussed herein appear to have come fromrelatively mature theropods based on the complete closure of theneurocentral sutures (Brochu, 1996; Irmis, 2007). The nomencla-ture for vertebral laminae and fossae used herein follows thatoutlined by Wilson (1999) and Wilson et al. (2011).

3. T’an’s Site, April 1923

T’an’s site, from which the four theropod vertebrae describedbelow were recovered, was located w500 m (1 li) southwest of avillage called Chiang Chün Ting, also known as Tianqiaotun orT’ien-Ch’iao-T’un (Buffetaut, 1995). Based on the labels on thespecimens in the PMU, the holotype specimen of Tanius sinensiswas found at the same locality, as were some of the ankylosaurspecimens described by Buffetaut (1995).

3.1. PMU 24711 (Cervical Vertebra)

PMU 24711 was mentioned as a vertebra of indeterminate po-sition, but not described or illustrated, by Wiman (1929). Theposterior portion of the centrum, the postzygapophyses, the base ofthe neural arch, the partial right prezygapophysis and the dorsalsurface of the right diapophysis are preserved. The right para-pophysis may also be preserved; if so, it remains encased in matrix(Fig. 2). The bone is extremely fragile: in several places, the corticalbone has peeled away, revealing the internal pneumatisation of thevertebra, which consists of large holes separated by thin walls ofbone.

The posterior articular facet of the centrum, which appears tobe completely preserved, is flat, suggesting that the vertebra wasamphiplatyan. In lateral view, the posterior articular facet isangled anterodorsally/posteroventrally relative to the flat ventralmargin of the centrum. The lateral surface of the posteriorcentrum is shallowly concave. As far as can be observed, this fossa(which does not house a foramen) was bounded dorsally by thebase of the subtly expressed posterior centrodiapophyseal lamina(PCDL).

The centropostzygapophyseal laminae (CPOLs) form the lateralmargins of the posterior neural canal. The gap between theventrolaterally facing postzygapophyses is bridged by a thin

intrapostzygapophyseal lamina (TPOL) which dips ventrally at themid-line. A bony strut, presumably a hyposphene, was attached tothe ventral surface of this lamina at the mid-line, though it isincompletely preserved and partially obscured by matrix. Themorphology of the medial surfaces of the lateral walls of the neuralcanal suggests that thin struts of bone ascended towards the mid-line of the neural arch, meeting the hyposphene and lending itstructural support. These struts would have formed the dorsalmargin of the neural canal, and also created paired cen-tropostzygapophyseal fossae (CPOFs) located dorsolateral withrespect to the neural canal. The dorsal surface of the TPOL ismedially concave, forming a spinopostzygapophyseal fossa (SPOF)which is accentuated by the stout spinopostzygapophyseal laminae(SPOLs) which connect the postzygapophyses to the mediallylocated neural spine. A horizontal postzygodiapophyseal lamina(PODL) connects the right postzygapophysis to the right dia-pophysis, and a prezygodiapophyseal lamina (PRDL) appears tohave present between the diapophysis and the right pre-zygapophysis. The diapophysis projects ventrolaterally from theanterior third of the vertebra, though only part of its dorsal surfacecan be observed; both it and the parapophysis are possibly pre-served but, if so, remain embedded in sediment.

3.1.1. PMU 24711 ComparisonsWithin Theropoda, amphiplatyan cervical vertebrae are

commonly observed in Coelurosauria (Gauthier, 1986; Holtz, 2000;Norell et al., 2001; Turner et al., 2012), though there are excep-tions. The cervicals of Compsognathus longipes Wagner, 1861 wereinterpreted as opisthocoelous by Ostrom (1978) in the German typespecimen, but later suggested to be platycoelous like those of Sino-sauropteryx prima Ji and Ji,1996 by Currie and Chen (2001). However,Peyer (2006) found that they were opisthocoelous in the Frenchreferred specimen (type specimen of Compsognathus corallestrisBidar, Demay, and Thomel, 1972). Eotyrannus lengi Hutt, Naish,Martill, Barker, and Newbery, 2001, Raptorex kriegsteini Sereno, Tan,Brusatte, Kriegstein, Zhao, and Cloward, 2009, and Dilong para-doxus Xu, Norell, Kuang, Wang, Zhao, and Jia, 2004 are some of thefew other coelurosaurs to have opisthocoelous cervical vertebrae(Benson, 2008), whereas some tyrannosaurids, like Tyrannosaurusrex Osborn, 1905 and Alioramus altai Brusatte, Carr, Erickson, Bever,and Norell, 2009, retained (or, perhaps, redeveloped) slightly opis-thocoelous cervical vertebrae as well (Brochu, 2003; Brusatte et al.,2012). The only maniraptoran theropods known to have opistho-coelous cervical vertebrae are alvarezsaurus (Turner et al., 2012).

Bevelled amphiplatyan centra (i.e. centra with articular facesthat are angled relative to their ventral horizontal long axis) arepresent in oviraptorids and possibly caenagnathids (Balanoff and

Fig. 2. Theropod cervical vertebra, PMU 24711. A, left lateral view; B, posterior view; C, dorsal view; D, right lateral view; E, ventral view. Scale bar ¼ 50 mm.

S.F. Poropat, B.P. Kear / Cretaceous Research 45 (2013) 103e113106

Norell, 2012), as well as in paravians (Turner et al., 2012) likeSinornithoides youngi Russell and Dong, 1993, Anchiornis huxleyi Xu,Zhao, Norell, Sullivan, Hone, Erickson, Wang, Han, and Guo, 2009and Deinonychus antirrhopus Ostrom, 1969a (Hu et al., 2009;Ostrom, 1969b; Russell and Dong, 1993). The cervical vertebrae ofthe tyrannosauroid Xiongguanlong baimoensis Li, Norell, Gao, Smith,and Makovicky, 2010 and the basal coelurosaur Coelurus fragilisMarsh, 1879 show similar bevelling of the articular facets(Carpenter et al., 2005; Marsh, 1881; Ostrom, 1980), but those ofCoelurus are muchmore elongate (Carpenter et al., 2005) than PMU24711; the cervical vertebrae of Xiongguanlong, on the other hand,more closely match the morphology of PMU 24711 (Li et al., 2010).

The centrum of PMU 24711 is quite short, suggesting that it doesnot derive from an ornithomimosaur; ornithomimosaur vertebraeare generally elongate and low (Makovicky et al., 2004). Theapparent presence of a hyposphene suggests that this vertebrapertains to a tyrannosauroid, since tyrannosauroids are the onlycoelurosaurs known to have hyposphenes on their cervical verte-brae (Makovicky, 1995). The descriptions of the anterior cervicalvertebrae of Tarbosaurus bataar (Maleev, 1955a) Maleev, 1955b arenot inconsistent with the morphology of this cervical vertebra;however, the vertebrae figured by Maleev (1974) are taller dorso-ventrally and shorter anteroposteriorly. This probably represents a

subadult feature of this vertebra (since tyrannosaur cervicals areknown to be anteroposteriorly compressed in adults and less so injuveniles (Holtz, 2004)); thus, it seems likely that PMU 24711represents a subadult tyrannosauroid, possibly Tarbosaurus bataar.

3.2. PMU 24712 (Cervical Vertebra)

PMU 24712 was identified as a cervical vertebra, brieflydescribed andmeasured, but not illustrated, byWiman (1929). Thisvertebra is much more complete than PMU 24711, lacking only theposterior cotyle, the anterior-most portion of the right pre-zygapophysis, and the cervical ribs (Fig. 3).

PMU 24712 is solid and blocky. The articular surfaces arebevelled very slightly relative to the centrum; this feature,coupled with the robusticity of the vertebra, suggest that PMU24712 is a more posterior cervical than PMU 24711. The anteriorarticular surface of the centrum is concave, whereas the posteriorarticular facet is incompletely preserved: it is probable that thevertebra was platycoelous in life. Wiman (1929) suggested thatthe vertebra might have been procoelous if the posterior surfaceconsisted of cartilage. However, this is unlikely, since no knownnon-avian theropods possess procoelous cervical vertebrae; thecervical vertebrae of Shanshanosaurus huoyanshanensis Dong,

Fig. 3. Theropod cervical vertebra, PMU 24712. A, anterior view; B, dorsal view; C, left lateral view; D, ventral view; E, posterior view; F, right lateral view. Scale bar ¼ 50 mm.

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1977 (¼ Tarbosaurus bataar) were interpreted as procoelous byHoltz (2001) but were shown to be platycoelous by Currie andDong (2001).

The ventral surface of the centrum is narrowand seems to bear asubtle mid-line keel. The parapophysis is located at the ante-roventral margin of the centrum. A very shallow fossa can be seenon the lateral surface of the centrum, bounded ventrally by theweakly expressed posterior centroparapophyseal lamina (PCPL)and dorsally by the PCDL. A raised ridge of bone is present imme-diately posterodorsal of the parapophysis, which separates themain fossa from a much smaller, shallower concavity on the dorsalmargin of the parapophysis.

The neural canal is subcircular in anterior view. Its ventralmargin corresponds to a slight depression in the dorsal margin ofthe anterior articular facet, and its ventrolateral margins areformed by stout centroprezygapophyseal laminae (CPRLs). TheCPRLs project dorsolaterally from the dorsal margin of thecentrum, and small accessory laminae emerging dorsomediallyfrom them form the dorsolateral margins of the neural canal.These accessory laminae do not meet on the mid-line, insteadbeing truncated by a thin, horizontal lamina which could beinterpreted as a very weak interprezygapophyseal lamina (TPRL).The neural canal could not be observed in posterior view as it isobscured by matrix.

The prezygapophyses are stout and widely spaced. Their artic-ular facets face dorsomedially to receive the more closely spaced,ventrolaterally facing postzygapophyses of the preceding vertebra.The prezygapophyses are connected to the neural spine via poorly-defined spinoprezygapophyseal laminae (SPRLs). The diapophysesare almost in direct contact with their respective prezygapophyses,though the intermediate regions appear to correspond to pre-zygodiapophyseal laminae (PRDLs). However, this region of thevertebra is poorly preserved on both sides, rendering confirmationof this interpretation difficult. Two of the most clearly preservedlaminae on the vertebra attach to the diapophysis: the sub-horizontal postzygodiapophyseal lamina (PODL) and the ante-rodorsallyeposteroventrally inclined PCDL.

A deep centrodiapophyseal fossa (CDF) is present on the ventralsurface of the PCDL and the base of the diapophysis, whereas ashallower postzygapophyseal centrodiapophyseal fossa (POCDF),open posterolaterally, is bounded dorsally by the PODL and ante-roventrally by the PCDL.

The neural spine is centred slightly posterior of the mid-lengthof the vertebra. It was clearly unbifurcated and was connectedto all four zygapophyses by poorly-defined laminae. The post-zygapophyses, which have reduced epipophyses, only meet at thebase of the neural spine, whereas the TPRLs which connect theprezygapophyses form a distinct shelf between the SPRLs imme-diately anterior of the neural spine. Based on the preservation ofthis element, it seems likely that the anterior face of the neuralspine was slightly concave; however, since this area is obscured bymatrix, it is not possible to rule out that it was actually flat.

3.2.1. PMU 24712 ComparisonsOverall, PMU 24712 appears most similar to the posterior-most

cervical vertebrae of tyrannosaurs and ornithomimosaurs. Orni-thomimosaur posterior cervicals are platycoelous (Makovicky,1995), as in Struthiomimus altus (Lambe, 1902) Osborn, 1917,Archaeornithomimus asiaticus (Gilmore, 1933) Russell, 1972,Gallimimus bullatus Osmólska, Roniewicz, and Barsbold, 1972and Pelecanimimus polyodon Pérez-Moreno, Sanz, Buscalioni,Moratalla, Ortega, and Rasskin-Gutman, 1994 (Gilmore, 1933;Osborn, 1917; Osmólska et al., 1972; Pérez-Moreno et al., 1994);the posterior-most cervical of A. asiaticus was reported to havebeen opisthocoelous (Smith and Galton,1990), though this appearsto be an exception among ornithomimosaurs. The posterior-mostcervicals of ornithomimosaurs also bear a mid-line keel(Makovicky, 1995), reduced epipophyses (Barsbold, 1983), andhave low neural spines (Osmólska et al., 1972); PMU 24712 has allof these features, further supporting this interpretation. Thisvertebra also shows similarities, including short, low neuralspines, reduced epipophyses, and overall relative proportions, tothe posterior cervicals of some tyrannosauroids, such as Tarbo-saurus bataar and Alioramus altai (Brusatte et al., 2012; Maleev,

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1974). However, the zygapophyses in A. altai and T. bataar aremuchmore widely spaced than in PMU 24712; in the latter case, this isdespite the fact that the vertebra is significantly smaller (A. altaicentrum length 75 mm; PMU 24712 centrum length 119 mm). Thissuggests that PMU 24712 is more likely to be an ornithomimosaurthan a tyrannosauroid.

3.3. PMU 24713 (Dorsal Vertebra)

PMU 24713 is almost complete, lacking only the tips of thezygapophyses and parapophyses and the right diapophysis (Fig. 4).Based on the position of the parapophysis on the neural arch, thisvertebra is a posterior dorsal. The bone appears to have experi-enced some slight transverse compression, as shown by thedifferent positions of the pneumatic foramina on each side.

The centrum is platycoelous, with the posterior face somewhatshallower. This contrasts with the “prosulcate” (i.e. with an anteriorconcave groove rather than a pit), dorsal vertebrae of Tyrannosaurusrex (Brochu, 2003). The centrum is spool-shaped, being constrictedat mid-length but flaring at both ends. The ventral surface of thecentrum appears to have borne a mid-line keel in life, thoughdeformation has distorted and exaggerated this feature. Smallpneumatic foramina, which are not set in fossae, are present onboth sides, with the right foramen situated lower than the leftrelative to centrum height. Based on the morphology of theforamina, we interpret that the right foramen has experienced lessdeformation and more closely reflects the original morphology: asmall, yet deep, foramen penetrating into the lateral surface of thecentrum.

Fig. 4. Theropod dorsal vertebra, PMU 24713. A, anterior view; B, dorsal view; C, left late

The neural arch is almost twice as tall as the centrum (Table 2),and is highly pneumatised. Viewed anteriorly, the neural canal isoval and transversely compressed, bound laterally by CPRLs butopen dorsally to approximately one-quarter of the length of thevertebra due to the absence of a TPRL. Thus, between the pre-zygapophyses, a deep cleft is present (the hypantrum) whichgradually shallows as it ascends towards the base of the neuralspine. The prezygapophyses faced dorsomedially, and were con-nected to the neural spine by weakly expressed SPRLs.

The bases of the parapophyses lie on the laminae interpretedabove as CPRLs, thus preventing the development of anterior cen-troparapophyseal laminae (ACPLs); however, a case could be madefor the part of the CPRL ventral to the parapophysis to be interpretedas the ACPL, with the dorsal portion (which joins the parapophysisto the prezygapophysis) becoming the prezygodiapophyseal lamina(PRDL). However, given that the medial surface of the laminaextending between the prezygapophysis and the centrum iscontinuous on its medial surface, it seemsmore sensible to interpretit as a single lamina.

In lateral view, the parapophysis bridges the anterodorsal part ofthe centrum and the anteroventral portion of the neural arch. Theparapophysis is located anteroventral relative to diapophysis and isnot connected to it by a parapodiapophyseal lamina (PPDL).Immediately posterior of the parapophysis an anterodorsallyeposteroventrally angled lamina is present which may represent thePCPL. This lamina has been ventrally truncated by a feature inter-preted herein as the anterior centrodiapophyseal lamina (ACDL).The coalescence of the bases of these two laminae has resulted inmuch of the lateral surface of the neural arch ventral to their

ral view; D, posterior view; E, right lateral view; F, ventral view. Scale bar ¼ 50 mm.

Table 2Measurements of the theropod vertebrae held in the PMU from the Upper Cretaceous Wangshi Group of Shandong Province, eastern China.

Measurements(mm)

Overallheight

Centrum Length from prezygapophysisto postzygapophysis

Neuralarch height

Neural canal

Maximumlength

Anterior Posterior Anterior Posterior

Height Width Height Width Height Width Height Width

PMU 24711 64 e e e 40 35 104 e e e 20 18PMU 24712 100 119 50 43 47 30 127 50 18 11 e e

PMU 24713 190 60 66 48 65 52 84 124 25 8 23 9PMU 24714 >21 48 12 17 12 18 e e e e e e

PMU 24719 >30 46 22 29 21 27 >36 e 4 6 4 5

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intersection being smooth. The PCPL, ACDL and the pre-zygodiapophyseal lamina (PRDL) enclose a ventrally pointed,triangular fossa which seems to be a prezygapophyseal cen-trodiapophyseal fossa (PRCDF); a similar fossa can be observed inthe anterior dorsal vertebrae, specifically dorsals 2e4 (Holtz, 2004),of Tarbosaurus bataar (Maleev, 1974).

The diapophysis is angled strongly dorsolaterally, but is notswept anteriorly or posteriorly. It is connected directly to the pre-zygapophysis by a thin PRDL and supported from below by theanteroventrally truncated ACDL and a more completely andstrongly expressed PCDL. The intersection between the dorsalsurface of the diapophysis and the lateral surface of the neuralspine bears a small, shallow fossa. A much more extensive, deeperfossa (the postzygapophyseal centrodiapophyseal fossa [POCDF]),enclosed by the PCDL, centropostzygapophyseal lamina (CPOL), andPODL, is located on the posterior surface of the diapophysis andextends dorsally on the ventral surface of the PODL, forming a deepconcavity.

The postzygapophysis is supported ventrally by the CPOL,attached to the diapophysis by a PODL and to the neural spine by aspinopostzygapophyseal lamina (SPOL). A deep sulcus is presentbetween the paired SPOLs on the posterior surface of neural spine,though this constricts posteroventrally as it approaches the bases ofthe postzygapophyses.

The posterior opening of the neural canal is laterally borderedon each side by the shared base of the PCDL and CPOL. The CPOLswrap around the neural canal, forming its roof as they meet at themid-line to form an undivided hyposphene, then diverging tosupport the paired postzygapophyses.

Fig. 5. Theropod caudal vertebra, PMU 24714. A, posterior view; B, dorsal view; C, right la

The neural spine is unbifurcated, oriented vertically, and centredon the posterior third of the centrum. The SPOLs overhang theposterior margin of the centrum. Roughened scars for ligamentattachment can be observed on the anterior and posterior faces ofthe neural spine.

This dorsal vertebra broadly resembles an isolated (and 1/3larger) theropod dorsal reported by Young (1958) from HsikouQuarry, Laiyang, Shandong.

3.3.1. PMU 24713 ComparisonsThe undivided hyposphene characterises PMU 24713 as a

non-maniraptoran theropod (Turner et al., 2012). In overallmorphology, PMU 24713 corresponds well to presacral 21 ofTyrannosaurus rex, especially with respect to the orientation of thediapophyses (Brochu, 2003). Superficially, it is not dissimilar fromthe dorsal vertebrae of ornithomimosaurs like Garudimimus bre-vipes Barsbold, 1981 and Gallimimus bullatus; however, the neuralarch in these taxa are seldom one-and-a-half times taller than thecentrum (Kobayashi and Barsbold, 2005; Osmólska et al., 1972),whereas in PMU 24713 the neural arch is almost double the heightof the centrum (Table 2). Based on this feature alone, it would seemthat PMU 24713 could be identified as a tyrannosauroid vertebra.

The brief description of the vertebrae of Yutyrannus huali Xu,Wang, Zhang, Ma, Xing, Sullivan, Hu, Cheng, and Wang, 2012stated that the vertebrae of this non-tyrannosaurid tyrannosau-roid were less pneumatised than those of Tyrannosauridae; thislends support to the identification of PMU 24713 as a subadulttyrannosaurid. Indeed, the vertebra closely matches the de-scriptions and images of the dorsal vertebrae of Tarbosaurus bataar

teral view; D, ventral view; E, anterior view; F, left lateral view. Scale bar ¼ 10 mm.

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provided by Maleev (1974): the centra are amphicoelous and tallerthan wide; weakly-developed, unseparated hyposphenesehypan-tra (mislabelled as zygosphenes-zygantra) are present; the dia-pophyses are supported from below by thin, plate-like PCDLs; andthe neural spine is transversely narrow and anteroposteriorly long.Thus, it is probable that PMU 24713 pertains to a subadult tyran-nosaurid, possibly Tarbosaurus bataar.

3.4. PMU 24714 (Caudal Vertebra)

PMU 24714 is represented by a centrumwith partial neural archand a broken prezygapophysis (Fig. 5). The centrum is elongate,dorsoventrally compressed, and weakly constricted at mid-length(Table 2). The ventral surface is slightly concave both ante-roposteriorly and transversely, bounded laterally by subtle, ante-roposteriorly extending ridges. The articular faces, though worn,suggest that the centrum was platycoelous in life. In anterior andposterior views, the centrum appears sub-hexagonal due to thedivision of the lateral surface into a lower ventrolaterally facingsurface and an upper dorsolaterally facing surface. Posteriorly, theupper surface encroaches on the lower. Transverse processes areabsent. The above observations, coupled with the relatively smallsize of the vertebra (Table 2), this suggests that this is a distal caudal.

The bases of the widely spaced prezygapophyses are preserved,whereas no sign of the postzygapophyses is present. The neuralcanal is small, and dorsoventrally compressed.

3.4.1. PMU 24714 ComparisonsThe incompleteness of both articular ends, and the fact that the

majority of the neural arch is missing, make precise assessment of

Fig. 6. Theropod caudal vertebra, PMU 24719. A, posterior view; B, dorsal view; C, right la

the affinities of this specimen difficult. Dorsoventrally compressedcentra with ventral median sulci bordered laterally by ridges arecharacteristics of ornithomimids (Longrich, 2008; Makovicky,1995), suggesting that PMU 24714 derives from an ornithomimid.Ornithomimids with distal caudal vertebrae showing these featuresinclude the un-named “Huren-duh (or Khuren-dukh) ornithomi-mosaur” (GIN 960910KD), Gallimimus bullatus, Ornithomimusedmontonicus Sternberg, 1933, and an un-named ornithomimidfrom Tajikistan (PIN 3041/12 and 3041/13) (Alifanov and Averianov,2006; Kobayashi, 2004; Makovicky, 1995; Osmólska et al., 1972).

4. Zdansky’s Site, October 1923

Zdansky’s site yielded only two specimens representing thero-pods: a caudal vertebra and an ungual phalanx. No other dinosaurremains in the collection are known to have come from this site(Wiman, 1929).

4.1. PMU 24719 (Caudal Vertebra)

PMU 24719 is longitudinally shorter, dorsoventrally taller andmore robust than PMU 24714 (Table 2). The centrum is virtuallycomplete, though the articular facets areworn. The neural arch onlypreserves the bases of the zygapophyses; the neural spine appearsto have been reduced, and there is no base for a transverse processpreserved, suggesting that this vertebra is a middle or distal caudal,either within or posterior of the “transition point” sensu Russell(1972).

The centrum isplatycoelous, spool-shaped, seeminglycompletelyapneumatic, and sub-round in anterior and posterior views (Fig. 6).

teral view; D, ventral view; E, anterior view; F, left lateral view. Scale bar ¼ 20 mm.

Table 3Measurements of the manual ungual phalanx from theWangshi Group of ShandongProvince, China, held in the PMU.

Measurements(mm)

Length Proximal Groovelength

Dorsoventral Mediolateral Left Right

PMU 24717 37 19 7 21 23

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Lateral ridges on the centrum split the lateral surface into dorsolat-erally and ventrolaterally facing portions (as in PMU 24714), how-ever, despite the fact that the ridges flare at the posterior end, thecentrum does not appear hexagonal in anterior or posterior viewbecause of the rim surrounding each articular facet. The ventralsurface of the centrum bears a narrow sulcus flanked by two ridges;this is most strongly expressed at the mid-length but weakens to-wards the articular surfaces.

The neural canal is dorsoventrally compressed. The neural archpreserves distinct, concave upwards semicircular ridges on itslateral faces which delineate the bases of the zygapophyses: thelower ridge on each side leads to the postzygapophysis, the upperto the prezygapophysis. The dorsal surface of the upper ridge issmooth and anteroposteriorly concave, with the base of thereduced neural spine and the intersection of the postzygapophysesthe only features interrupting its surface. In dorsal view, a thin ridgeextends posteriorly from each prezygapophysis; these ridges unitejust posterior of the mid-length of the vertebra. The pre-zygapophyses would have been further apart than the post-zygapophyses, though no other details of their morphology can bestated with certainty due to their incomplete preservation.

4.1.1. PMU 24719 ComparisonsPMU 24719 shares features with the caudal vertebrae of several

coelurosaur groups. The presence of pleurocoels in all but thedistal-most caudal vertebrae of Citipati osmolskae Clark, Norell andBarsbold, 2001 suggests that PMU 24719 does not pertain to anoviraptorosaur (Balanoff and Norell, 2012). Troodontid caudalvertebrae are too poorly known to allow comprehensive compari-sons to be made (Makovicky and Norell, 2004); in any case, thosethat are well known tend to be much more elongate than PMU24719 (Russell and Dong, 1993). The neural spines of most orni-thomimosaur and tyrannosauroid caudal vertebrae tend to be

Fig. 7. Theropod ?right ungual phalanx, PMU 24717. A, dorsal view; B, medial v

centred towards the posterior of the centrum, as in PMU 24719(Brochu, 2003; Osmólska et al., 1972); however, given that thisvertebra does not have an elongate centrum, yet is past the tran-sition point based on the absence of transverse processes, it is likelyto represent a tyrannosauroid (Makovicky, 1995).

4.2. PMU 24717 (Manual Ungual Phalanx)

PMU 24717 is a small ungual phalanx, which we interpret as amanual claw due to it being transversely (rather than dorsoven-trally) compressed (Table 3). The claw is incomplete at the tip andalso at its proximodorsal margin (Fig. 7). The claw is gently curved,and the flexor tubercle is prominent, elongate and bulges slightlymore to the left side of the claw than the right. The blood vesselgrooves extend from the tip of the claw only as far back as the flexortubercle.

4.2.1. PMU 24717 ComparisonsThe extents of the blood vessel grooves, the development and

morphology of the flexor tubercle, and the degree of curvature ofPMU 24717 correspond well with manual ungual I of the holotypespecimen (BSPAS I 563) of Compsognathus longipes, and themanualunguals of Gallimimus bullatus (Gishlick and Gauthier, 2007;Osmólska et al., 1972; Ostrom, 1978). The blood vessel groovesare unlike those of deinonychosaurs which extend along the wholelength of the claw from the articular facet to the tip (Lü et al., 2010;Ostrom, 1969b; Russell and Dong, 1993).

5. Summary

Despite the fact that the Wangshi Group has produced fewtheropods to date, the material that it has yielded offers a tanta-lising glimpse at what appears to have been a coelurosaur-dominated theropod fauna living in eastern China during the LateCretaceous. Non-coelurosaurian theropods dominate the upper-most Cretaceous rocks of Asia and North America (Weishampelet al., 2004), rendering this finding unsurprising. Furthermore,the majority of these specimens, appear to pertain to tyrannosau-roids: it is probable, based on their age and their favourable com-parison with specimens of Tarbosaurus bataar, that they representsubadult tyrannosaurids. These remains, though fragmentary,suggest that future excavations in the Wangshi Group, Shandong

iew; C, ventral view; D, proximal view; E, lateral view. Scale bar ¼ 10 mm.

S.F. Poropat, B.P. Kear / Cretaceous Research 45 (2013) 103e113112

Province will reveal more about the theropods which lived along-side the much more common and better represented large hadro-saurs in China during the Late Cretaceous.

Acknowledgements

The authors would like to warmly thank: J. O. R. Ebbestad(Museum of Evolution, Uppsala, Sweden) for allowing us access tospecimens in his care; P. Eriksson (Museum of Evolution, Uppsala,Sweden) for his skilful repair of the specimens; and M. Mortimerand P. J. Makovicky (Field Museum of Natural History, Chicago) forassistance in obtaining articles cited in this work. We would alsolike to acknowledge S. L. Brusatte (University of Edinburgh, Scot-land) and an anonymous reviewer for their helpful critique of themanuscript. Finally, we acknowledge the Australian ResearchCouncil for funding for this work through ARC Grant LP100100339,awarded to B. P. Kear as Chief Investigator.

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