Historical Biology, 2014 Vol. 26, No. 2, 236251, http://dx.doi.org/10.1080/08912963.2013.

809347

An early juvenile specimen of Bolong yixianensis (Ornithopoda: Iguanodontia) from the Lower Cretaceous of Ningcheng County, Nei Mongol, China
Wenjie Zhenga*, Xingsheng Jina, Masateru Shibatab and Yoichi Azumaa,b
a

Zhejiang Museum of Natural History, Zhejiang 310014, P.R. China; bFukui Prefectural Dinosaur Museum, Fukui 911-8601, Japan

(Received 2 March 2013; nal version received 21 May 2013; rst published online 1 July 2013) We describe an early juvenile specimen (ZMNH M8812) of Bolong yixianensis from the Yixian Formation (Lower Cretaceous) of Ningcheng County, Nei Mongol, China. The specimen consists of an almost complete skeleton preserved two-dimensionally on a slab. The short and deep skull proportions and unfused neurocentral sutures in most preserved vertebrae suggest that the ZMNH M8812 is a juvenile individual. Osteohistological study conrms a very early developmental stage. The study reveals the ontogenetic changes of Bolong for the rst time. The specimen revealed one additional autapomorphy for Bolong yixianensis: the lingual face of the maxillary crown is bounded by thickened mesial and distal margins and bisected by a prominent median principal ridge. The study revealed the following ontogenetic trends of Bolong: increased tooth rows in both maxilla and dentary, increased robustness of the jugal and scapula, the radius and ulna become more robust and shorter relative to the hindlimb and the metatarsals become proportionally shorter. ZMNH M8812 represents the rst juvenile non-hadrosaurid iguanodontian specimen described from the Lower Cretaceous of eastern Asia. Keywords: Bolong yixianensis; juvenile; Iguanodontia; Lower Cretaceous; Yixian Formation; China

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

Introduction Complete ontogenetic series currently exist for several Hadrosaurid dinosaurs (Horner and Currie 1994; Evans rquez 2007; Brink et al. 2010; Evans 2010; Prieto-Ma 2011). In contrast, the fossil material available for most non-hadrosaurid iguanodontians is insufcient for documenting the full range of morphological changes through rquez 2011). Nevertheless, several ontogeny (Prieto-Ma specimens of juvenile non-hadrosaurid iguanodontians have contributed to ontogenetic studies (Forster 1990a, bner and Rauhut 2010; Werning 2012). 1990b; Hu However, juvenile individuals, especially in early stages of development, and ontogenetic series of skeletal elements of non-hadrosauroid iguanodontians remain relatively rare. Although iguanodontian dinosaurs are very common in east-central Asia (Norman 1996, 1998, 2002; Wang and Xu 2001; You et al. 2003, 2005), no juvenile non-hadrosaurid iguanodontian from Asia has yet been described. Barrett et al. (2009) mentioned that several specimens possibly referable to Jinzhousaurus were excavated from Yixian County of Liaoning Province (Figure 1), but none of these have been described. Here, we describe an early juvenile specimen of Bolong yixianensis from Xitaizhi Village, Shantou Town, Ningcheng County, Nei Mongol Autonomous Region, Peoples Republic of China (Figure 1). The specimen was collected by a local villager, and detailed information of the fossil locality remains unclear. It most probably

belongs to the lower part of Yixian Formation (Wang et al. 2000, 2005; Liu et al. 2006). The specimen was prepared by Yuqing Zhang, under the supervision of Dr Junchang in the Institute of Geology, Chinese Academy of Lu conrmed authenGeological Sciences, Beijing. Dr Lu ticity of the specimen.

Systematic paleontology Dinosauria Owen, 1842 Ornithischia Seeley, 1887 Ornithopoda Marsh, 1881 Iguanodontia Dollo, 1888 Bolong yixianensis Wu, Godefroit and Hu, 2010 (Figures 2 15, Table 1) Material The specimen is a nearly complete skeleton housed in Zhejiang Museum of Natural History, Hangzhou, China (ZMNH M8812) (Figures 2 15, Table 1).

Locality and horizon Xitaizhi Village, Shantou Town, Ningcheng County, Nei Mongol Autonomous Region, China. Lower Cretaceous, the lower part of Yixian Formation.

*Corresponding author. Email: zhengwenjie@gmail.com This article was originally published with an error. This version has been amended. Please see Corrigendum (DOI: http://dx.doi.org/ 10.1080/08912963.2014.899182)
q 2013 Taylor & Francis

Historical Biology 237 similarities with the holotype of Bolong yixianensis. Both in Bolong and ZMNH M8812, the coronoid process is inclined caudodorsally, maxillary teeth possess one primary and several accessory ridges, dentary teeth possess a primary and a secondary ridge, the scapula is slender and straight with slight distal expansion, and ungual phalanx of digit I is large and conical (Wu and Godefroit 2012). Therefore, M8812 is most probably a juvenile Bolong yixianensis.

Estimation of the growth stage of ZMNH M8812 Abundant evidence suggests that this individual is very young. It is very tiny, with the whole body length , 50 cm. There are only eight vertical tooth positions (alveoli) preserved per maxilla and dentary, whearas there are at least 10 tooth positions in iguanodontians: 10 in the basal iguanodontian Zalmoxes (Weishampel et al. 2003), 15 in the maxilla and 14 in the dentary of the holotype of Bolong (Wu and Godefroit 2012), 16 in the dentary of the adult Jinzhousaurus (Barrett et al. 2009) and 23 in Probactrosaurus (Norman 2002). The tooth rows of both maxilla and dentary increase with the age of the individual in iguanodontians (Gilmore 1933; Weishampel et al. 2003; bner and Rauhut 2010). Open neurocentral sutures Hu occur in most preserved vertebrae, including the cervical, dorsal and anteriormost 17 caudal vertebrae. The condition in more posterior caudals is unclear due to poor preservation. In addition, the sacral centra are not fused with each other. The sequence of neurocentral suture closure is one criterion for ontogenetic stage determination in extant crocodylians, and was also used in dinosaurs (Brochu 1996; Irmis 2007), and ornithischian dinosaurs may have had a posterior-to-anterior progressing sequence of neurocentral suture closure (Irmis 2007). In case of this specimen, the open neurocentral sutures on at least the 17 anteriormost caudal vertebrae suggest a very young age at the time of death. To better assess the age of specimen ZMNH M8812 at the time of death, transverse mid-diaphyseal sections were taken from the right femur (Figure 3). The cortical bone consists of woven to poorly developed bro-lamellar bone tissue, which has a rather spongy appearance and consists of a network of bone trabeculae separated by large vascular canals. No line of arrested growth can be observed. There is hardly any centripetal deposition of primary osteonal material at the periphery of the canals. The microstructure of the bone is similar to that of Maiasaura bones at nestlings stage, and suggests that this individual was probably less than one year old when it died (Horner et al. 2000). The predominance of woven bone is also consistent with the still early ontogenetic stage of the individual, because the woven bone is considered to be the result of relatively rapid rates of bone deposition

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

Figure 1. Geographic location of ZMNH M8812 and holotype of Bolong (YZH-001).

Taxonomic identication of ZMNH M8812 ZMNH M8812 can be assigned to Iguanodontia based on the following synapomorphies of Iguanodontia: a deep dentary ramus with parallel dorsal and ventral borders, and compressed and blade-shaped prepubic process (Norman 2004). The conical ungual phalanx of manus digit I suggests that ZMNH M8812 is a basal (non-hadrosaurid) iguanodontian (Norman 2004). Iguanodontians are very common in north China, Mongolia and Russia (Weishampel et al. 2004). Among the basal iguanodontians found from central Asia, ZMNH M8812 is most similar to Bolong yixianensis. Jinzhousaurus (Wang and Xu 2001) and Bolong (Wu et al. 2010) are also known from the Yixian Formation. ZMNH M8812 can be distinguishable from the holotype of Jinzhousaurus yangi by several cranial and postcranial features: the coronoid process of Jinzhousaurus is perpendicular with long axis of the dentary, whereas the coronoid process is inclined caudodorsally in ZMNH M8812. The prominent primary ridge in labial surface of Jinzhousaurus is stronger than that in ZMNH M8812 (Barrett et al. 2009). The scapular shaft in Jinzhousaurus is broader than that in M8812, the distal expansion of scapula in Jinzhousaurus is also stronger (Wang et al. 2010). ZMNH M8812 shows many

238 W. Zheng et al.

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

Figure 2. ZMNH M8812. (A) Block containing nearly complete skeleton; (B) line drawing of A. Grey areas indicate cracks in the slab. Abbreviations: a, astragalus; c, calcaneum; cdv, caudal vertebra; ch, chevron; co, coracoid; cv, cervical vertebra; d, dentary; drb, dorsal rib; dv, dorsal vertebra; fem, femur; b, bula; hm, humerus; hy, hyoid; il, ilium; isc, ischium; j, jugal; mc, metacarpal; mt, metatarsal; mx, maxilla; n, nasal; pd, predentary; ph, phalanx; pm, premaxilla; po, postorbital; prf, prefrontal; pub, pubis; ra, radius; sa, surangular; sc, scapula; tib, tibia; ul, ulna.

(Cerda and Chinsamy 2012). The marrow cavity is indiscernible; this maybe the result of crushing of the bone due to sediment compaction. Alternatively, it may reect the very young ontogenetic stage of this individual.

Description Skull and mandible The skull appears to be essentially complete but disarticulated, the result of dorsoventral compression (Figures 4 and 5). The elements are crushed and overlay one another, and most components of skull can only be observed in medial or ventral view. In addition, most elements are badly damaged, restricting the amount of anatomical information that can be obtained from the specimen. The skull is relatively shorter and deeper than the adult Bolong (Wu and Godefroit 2012), and resembles those of basal ornithopods and basal iguanodontians (Weishampel et al. 2003). These proportions reect the

young stage of this individual, as in the other iguanodontians that the overall juvenile skull is proportionally shorter bner and and deeper than the adult skull (Evans 2010; Hu Rauhut 2010). Both premaxillae are disarticulated and exposed in medial view (Figures 2, 4 and 5). The nasal process is slender and gently arched dorsally. The maxillary process is robust in medial view and increases in dorsoventral depth posteriorly, similar to the adult Bolong (Wu and Godefroit 2012). The morphology of distal end of the maxillary process is unclear, due to the overlay of the right dentary. The anteroventral portion of the external naris, which is formed by the nasal and maxillary processes, is elongated and points anteroventrally. The anterior portion of external naris is slightly narrower and extends more anteriorly than that in the adult Bolong (Wu and Godefroit 2012). The rostral end of the premaxilla is strongly downturned like that in the adult Bolong, which also occurs in Jinzhousaurus, Altirhinus, Equijubus, Protohadros and hadrosaurids (Barrett et al. 2009; Wu and Godefroit 2012).

Historical Biology 239

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

Figure 3.

Osteohistology of the mid-diaphyseal femur of ZMNH M8812. (A) Cross section of mid-diaphyseal femur. (B, C) Details of A.

Both maxillae are present in medial view in the slab, although the anterior end is broken (Figures 2, 4 6). The right maxilla was prepared such that it can be observed in both medial and lateral views. Maxillae are anteroposteriorly elongate, with a low triangular outline, and their dorsal apex is positioned roughly at mid-length of the element. In the adult Bolong, the maxilla is relatively deeper and the apex is caudal to the centre of the maxilla (Wu and Godefroit 2012). The anteriormost ventral margin of the maxilla is edentulous. The right maxilla exhibits

Figure 4.

Photograph of skull and mandible of ZMNH M8812.

approximately eight vertical tooth positions, the maxilla in the adult has 10 tooth positions (Wu and Godefroit 2012). The ventral edge of tooth line is straight as in the adult. The thin alveolar parapet that covers the teeth is incompletely preserved, and no foramina dorsal to the parapet are observed. As in the adult Bolong, the anterior tip is less distinctly downturned than in Jinzhousaurus, Altirhinus, Equijubus, Shuangmiaosaurus and Protohadros (Wu and Godefroit 2012). Both nasals are present in ventral view (Figures 2, 4 and 5). The two nasals are sutured together for much of their length, but they diverge and taper anteriorly. In contrast, the nasals of the adult Bolong do not diverge anteriorly (Wu and Godefroit 2012). The jugal is well preserved in medial view (Figures 2, 4 and 5), and consists of three main processes: an anteriorly directed maxillary process, a posterodorsally directed postorbital process and a posteriorly situated quadratojugal process. The jugal in the holotype of Bolong is poorly preserved (Wu and Godefroit 2012). The ventral border of the jugal is sinuously curved. The maxillary process is sub-triangular in outline and tapers in height anteriorly; the medial surface of the maxillary process bears an anteroposteriorly elongated slot into which the jugal process of the maxilla would t. The maxillary process is more robust in the adult Bolong (Wu and Godefroit 2012). The postorbital process is anteroposteriorly narrow and tapers dorsally. The process curves slightly anteriorly. It is mediolaterally expanded, producing a sub-triangular to sub-rectangular transverse cross

240 W. Zheng et al.

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

Figure 5. Line drawing of skull and mandible of ZMNH M8812. Abbreviations: d, dentary; hy, hyoid; j, jugal; mx, maxilla; n, nasal; pd, predentary; pm, premaxilla; po, postorbital; prf, prefrontal; L, left; R, right.

section. It forms an angle of approximately 1008 with the maxillary process, the angle is smaller than that in the adult Bolong (approximately 1108) (Wu and Godefroit 2012). Posteriorly, the quadratojugal process is greatly expanded relative to the main body of the bone and has a dorsoventrally elongate posterior margin, which is concave in medial view. The body of the bone is deeper than that in Probactrosaurus (Norman 2002). The postorbital is a triradiate element consisting of three processes that converge to form a centrally positioned, sub-triangular main body (Figures 2, 4 and 5). These processes extend anteriorly, anteroventrally and posteriorly, respectively. It remains unclear whether the

anterior process contacts the prefrontal. The anteroventral (jugal) process is slender and well developed, similar to that seen in the majority of other basal iguanodontoids, whereas it differs from Equijubus, the postorbital of which lacks a well-developed ventral process (You et al. 2003; Barrett et al. 2009). The postorbital is poorly preserved in the holotype of Bolong (Wu and Godefroit 2012). Frontals are present in ventral view (Figures 2, 4 and 5). Near the midline, the posterior border of frontals unit is notched to form a V-shaped articular surface for the parietals. A similar feature is present in Jinzhousaurus, Altirhinus, but is absent in Dollodon, Iguanodon, Ouranosaurus and Probactrosaurus mazongshanensis

Figure 6.

Right maxilla of ZMNH M8812 in medial (A) and lateral (B) views.

Historical Biology 241

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

Figure 7. Select teeth of ZMNH M8812. Anterior teeth of right maxilla in lingual (A), occlusal (B) and labial (C) views; posterior teeth of right dentary in lingual (D) view. Abbreviations: os, occlusal surface; pr, primary ridge; r, ridge; sr, secondary ridge. Scale bar 5 cm.

(Barrett et al. 2009). The frontal is not preserved in the holotype of Bolong (Wu and Godefroit 2012). The partial predentary that supported the horny beak along its edentulous occlusal margin is preserved in

articulation with the left dentary (Figures 2, 4 and 5). The anterior margin of the predentary slopes anterodorsally towards its dorsal margin. The ventral process, articulate with the rostroventral border of the dentary, is well

Figure 8. Cervical vertebrae, dorsal ribs and pectoral girdle of ZMNH M8812. Abbreviations: acr, acromial process; co, coracoid; cv, cervical vertebra; drb, dorsal rib; dv, dorsal vertebra; gl, glenoid; sc, scapula; L, left; R, right. Scale bar 2 cm.

242 W. Zheng et al.

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

Figure 9. vertebra.

Dorsal vertebrae and ribs of ZMNH M8812. Abbreviations: drb, dorsal rib; dv, dorsal vertebra; il, ilium; pub, pubis; sv, sacral

developed, like the condition in the holotype of Bolong (Wu and Godefroit 2012). Both dentaries are exposed in medial view (Figures 2, 4 and 5). Their dorsal and ventral margins extend parallel to each other along the entire length of the tooth row. The rostral end of the dentary is slightly downturned as in the holotype of Bolong (Wu and Godefroit 2012). In Jinzhousaurus, Dollodon, Equijubus,

Fukuisaurus, Iguanodon, Lanzhousaurus, Mantellisaurus and Ouranosaurus, the anterior end of the dentary is also slightly downturned; a strongly deected dentary symphysis is present in Altirhinus, Protohadros and many hadrosaurids (Barrett et al. 2009). Anteriorly, the dentary contacts the predentary. The right dentary shows eight vertical tooth positions, fewer than that in the holotype of Bolong (Wu and Godefroit 2012). The teeth

Figure 10. The photograph (A, C) and line drawing (B, D) of the caudal vertebrae of ZMNH M8812. Abbreviations: cdr, caudal rib; ch, chevron; isc, ischium; mt, metatarsal; ns, neural spine. Scale bar 2 cm.

Historical Biology 243

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

Figure 11. The photograph (A) and line drawing (B) of humeri of ZMNH M8812. Abbreviations: dpc, deltopectoral crest; hm, humerus; L, left; R, right. Scale bar 2 cm.

are bordered medially by the alveolar parapet. The dentary tooth row is straight along its dorsal margin in medial view. There is a diastema between the posterior margin of the predentary and the rst dentary tooth. The coronoid process of right dentary is directed posterodorsally, identical to the holotype of Bolong (Wu and Godefroit 2012). This morphology is similar to that seen in Ratchasimasaurus (Shibata et al. 2011) and many nonankylopollexian ornithopods, such as Hypsilophodon (Galton 1974), Orodromeus (Scheetz 1999) and Dysalo-

bner and Rauhut 2010). Most of the derived tosaurus (Hu bner and iguanodontians possess an upright process (Hu Rauhut 2010). The Meckelian groove below the tooth alveoli extends along the whole length of the dentary, as in most non-ankylopollexian ornithopods and Camptosaurus dispar. In derived iguanodontians (including hadrosaurs) the Meckelian groove tapers and ends well bner and Rauhut 2010). posterior to the symphysis (Hu The overall morphology of dentary is very similar to that in the holotype of Bolong (Wu and Godefroit 2012); as

Figure 12. The photograph (A, C) and line drawing (B, D) of forearms and manii. Abbreviations: mc, metacarpal; ph, phalanx; ra, radius; ul, ulna; I V, digit numbers; 1 3, phalanx numbers; L, left; R, right. Scale bar 2 cm.

244 W. Zheng et al.

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

Figure 13. The photograph (A) and line drawing (B) of pelvic girdle of ZMNH M8812. Abbreviations: app, anterior pubic process; fem, femur; il, ilium; ipi, iliac peduncle of the ischium; isc, ischium; obt, obturator process; pp, posterior pubic process; ppi, pubic peduncle of the ischium; pub, pubis; L, left; R, right. Scale bar 2 cm.

in Hypacrosaurus, the embryonic dentary is similar in morphology to the corresponding adult element (Horner and Currie 1994).

Other parts of the skull, such as the surangular and hyoid, are also present; however, they provide no useful information.

Figure 14. Femur (A. B), tibia and bula (C, D) of ZMNH M8812. Abbreviations: fem, femur; b, bula; pub, pubis; tib, tibia; L, left; R, right. Scale bar 2 cm.

Historical Biology 245

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

Figure 15. Photograph (A, C) and line drawing (B, D) of pes ZMNH M8812. Abbreviations: a, astragalus; c, calcaneum; b, bula; mt, metatarsal; ph, phalanx; tib, tibia; tib; L, left; R, right. Scale bar 2 cm.

Dentition The maxillary dentition includes one active tooth per alveolus (Figures 4 7); each maxillary alveolus holds one active and one replacement tooth. The mesial and distal margins of the teeth bear strong denticles, which become smaller towards the apex. The shape of crown is mesiodistally wide, namely a leaf shape, rather than narrow. The enamel covers the entire lingual surface of the maxillary teeth, a condition that suggests it is a basal iguanodontian (Norman 2004). In the labial surface, the primary ridge is offset distally, similar to that in the holotype of Bolong (Wu and Godefroit 2012). The primary ridge is not prominent, weaker than that in the holotype of Bolong (Wu and Godefroit 2012). Up to six secondary ridges are developed, most of them are positioned mesial to the primary ridge. The secondary ridges are more developed than that in the holotype of Bolong (Wu and Godefroit 2012). Each ridge culminates in a denticle along the margin of the crown. The ridges are perpendicular to the wear facet, whereas in the holotype of Bolong, the primary ridge is deected distally (Wu and Godefroit 2012). The replacement crowns lie on the medial side of the roots of functional teeth. The lingual face of the crown is bounded by thickened mesial and distal margins and is bisected by a prominent median principal ridge, which is slightly offset distally. This character may represent an autapomorphy of Bolong. The lingual (cutting) edge of the occlusal surface displays a characteristic W-shaped view in occlusal view. As in all iguanodontians for which the dentition is known, the dentary teeth are arranged in a series of interlocking rows without spaces between the crowns. One active and one replacement tooth occurs in each alveolus, as in Probactrosaurus gobiensis (Norman 2002) and Bactrosaurus (Godefroit et al. 1998). The crowns of dentary teeth bear simple tongue-shaped denticles on the

mesial and distal margins (Figure 7(A)). The dentary crowns are broadly expanded anteroposteriorly and their overall proportion is close to the holotype of Bolong (Wu and Godefroit 2012), it also resembles those seen in Equijubus (You et al. 2003) and Altirhinus (Norman 1998). All dentary teeth are visible in lingual view. The enamelled lingual surface bears curved ridges running down from the mesial and distal margins that meet basally, enclosing a generally shield-shaped surface. The primary ridge is offset distally on the lingual surface and divides it into two asymmetrical areas. These areas are adorned by faint parallel accessory ridges that extend as buttresses from
Table 1. Selected measurements of ZMNH M8812 (in mm). Length Proximal width Minimum width Distal width Length Max. width Distal width Length Max. width Distal width Length Length Length Length Length Length Length Length Length Length Length Length Length Length 49.00 13.51 5.77 9.02 , 42.9 10.48 9.31 , 41.30 9.55 10.68 34.10 34.30 31.20 31.00 8.90 12.03 11.83 46.84 51.80 50.50 46.70 17.11 22.04 18.22

Right scapula

Right humerus Left humerus Right ulna Left ulna Right radius Left radius Left metacarpal II Left metacarpal III Left metcarpal IV Left ischium Left femur Left tibia Left bula Right metatarsal II Right metatarsal III Right metatarsal IV

246 W. Zheng et al. individual denticles on the crown margin in unworn crowns (Figure 7). The primary ridge is more prominent than on the maxillary teeth, contrary to the situation observed in the holotype of Bolong (Wu and Godefroit 2012). The primary ridge is more prominent than that in the holotype of Bolong (Wu and Godefroit 2012) and Probactrosaurus (Norman 2002). As in the adult Bolong, a secondary ridge is present mesial to the primary ridge, bisects the larger mesial half of the crown and reaches the upper part of the mesial margin; there may be a variable number of more or less parallel subsidiary ridges, which are extensions of the bases of the marginal denticles. The sacrodorsal and the sacrals are exposed in the lateroventral view (Figure 2). The sacrum and adjacent centra are unfused. There is no evidence of a ventral groove or ridge on the ventral surface of the sacral centra. The articular margins are expanded as in the preceding dorsals. The three anterior sacral ribs are articulated with the right ilium. Few other details of the sacral centra can be observed. The anterior and posterior series of caudal vertebrae are preserved in articulation, and a break occurs between the two sets (Figures 2 and 10). The articulated series are visible in the right lateral view. At least 17 anterior neural arches are unfused to their corresponding centra, whereas the condition in the posterior series of caudals is unclear. The 17 anterior caudal vertebrae are well preserved. In the anteriormost two caudals, the prezygapophyses project slightly beyond the anterior surface of the centrum. The neural arches are displaced from the centra in the more posterior elements of this series. Above the prezygapophyses, there is an elongated, slightly posteriorly inclined and transversely compressed neural spine. The lateral face of the centrum is slightly concave with the articular margins transversely expanded. The centra become longer and lower through the series. The caudal ribs are not fused to the centra. The centrum heights of the 12 most anterior caudals clearly exceed their lengths. In the more posterior ve caudals, the centra are slightly longer than high. The neural spines decrease in dorsoventral height along the series, and become increasingly posteriorly inclined. In the posterior series, the centra become more elongated, and the neural spines are more horizontally directed. Eleven chevrons are preserved in articulation with the caudal vertebrae (Figure 10). The posterior preserved chevrons are formed by two arches that join proximally to form the articular surface. The ventral rod tapers in both lateral and anterior views to terminate in a rounded tip. The chevrons are longer and more slender than the corresponding neural spine. The anterior six chevrons direct more caudally than the posterior chevrons. As in the holotype of Bolong (Wu and Godefroit 2012), the caudoventral orientation of posterior chevrons is apparently equivalent to the caudodorsal angle of the corresponding neural spine. The chevrons steadily become smaller along the series, forming a mirror image of the neural spines.

Vertebral column, ribs, chevrons The posterior cervical vertebrae form an articulated series (Figures 2 and 8). The cervicodorsal junction is obscured by the presence of the overlying right scapula and ribs (Figure 8). In lateral view, the main bodies of the centra are almost square in outline. The centra are deep and anteroposteriorly narrow. The centrum is compressed ventrally and forms a thin keel. Dorsal to this keel, the lower half of the centrum surface is concave, and develops a horizontal ridge on the upper half. The parapophysis is situated on the anterior end of this ridge. The anteriormost preserved rib was treated as the cervical rib. The rib is short bodied with the divergent rib head (Figures 2 and 8). The morphology of the cervical vertebrae and rib is generally similar to that of the holotype of Bolong (Wu and Godefroit 2012). Dorsal vertebrae are preserved in an articulated series (Figures 2 and 9). The anterior series are largely obscured by the overlying ribs and right scapula; the posterior dorsal vertebrae are exposed in the ventrolateral view. The centra are nearly equal in length. The ventral margin of the centra bears a longitudinal groove bounded by low longitudinal ridges. The centra are strongly expanded at the articular ends. The centra of dorsal vertebrae in holotype of Bolong are completely hidden by the scapula or the dorsal ribs, only the neural spines can be observed (Wu and Godefroit 2012); this condition hampers the comparison between the two specimens. The dorsal ribs are relatively short anteriorly (Figure 8), reach their maximal lengths in the anterior mid-trunk region and gradually decrease in size posteriorly (Figure 2). The posterior series of ribs are preserved in articulation with the vertebrae, and the articular ends are covered by the dorsal centra (Figure 9). The anterior series of ribs are double headed (Figure 8), with a larger capitulum, supported by a long capitular process, whereas the tuberculum forms only a very short articular process that extends in line with the rib shaft. The rib shaft is long and curved.

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

Pectoral girdle and forelimbs Both scapulae are present (Figures 2 and 8). The right element is complete and exposed in lateral view and it overlies the articulated series of the dorsal ribs. The left scapula is partially exposed in the medial view, whereas most of the elements are obscured by the overlying ribs. As in the holotype of Bolong, the proximal end is expanded dorsoventrally (distinctly deeper than the distal scapula) to

Historical Biology 247 support the glenoid and to provide a sutural surface for the coracoid, the acromial process is directed dorsally and the articular facet for the coracoid is extensive (Wu and Godefroit 2012). The scapular shaft is slightly curved along its length in the lateral view such that the dorsal margin of the blade is gently convex and the ventral margin is slightly concave. The curvature of the ventral margin is stronger than that of the dorsal margin. The posterior margin of the scapula is gently convex in the lateral view. The shaft is slightly slender than that in the holotype of Bolong, the ratio of scapular length to minimum width of the shaft is about 8.49, whereas the ratio is only 7.61 in the holotype of Bolong. Except this, the morphology of the scapula is generally similar to that of the holotype of Bolong (Wu and Godefroit 2012). Overall, the shaft is almost straight and only modestly expanded distally, similar to that of Probactrosaurus (Norman 2002), whereas the scapula of Jinzhousaurus is more strongly curved along its length and strongly expanded at the distal end (Wang et al. 2010). The right coracoid is present and is unfused to the scapula. In the lateral view, the anterodorsal margin is subcircular; the anteroventral corner of the coracoid forms a small hook-like process. Both humeri are preserved in the anterior view (Figures 2 and 11), and the humerus is approximately 81% of the length of the femur. Overall, the shaft of the humerus is slender, bowed and slightly sinuous in the anterior view. The proximal end is broad and has a shallowly concave anterior surface. The transverse width of the proximal expansion exceeds that of the distal end. The deltopectoral crest extends for approximately half of the length of the humerus. The deltopectoral crest curves anteriorly. The radial and ulnar condyles are separated by a shallow intercondylar groove. The humerus in the holotype of Bolong is also slender (Wu and Godefroit 2012). The proximal end of the ulna is anteroposteriorly expanded relative to the shaft, and the olecranon process is incomplete (Figure 12). The ulna is longer and more robust than the radius, The radius is slender, straight and anteroposteriorly expanded at either end to form the articular surfaces; it amounts , 74% of the length of the humerus. Compared to the holotype of Bolong (Wu and Godefroit 2012), the ulna and radius are more slender, the ratio of proximal width to length is only 0.15 in ulna and 0.19 in radius, much smaller than that in the holotype of Bolong (Wu and Godefroit 2012). Both manii are incomplete with the dorsal surface is extensively fractured (Figure 12). The elements of both hands are mainly articulated, but the distal phalanges of the left hand and the proximal elements of the right hand are missing. The left metacarpals II IV are closely appressed to one another and exposed in the dorsal view. Metacarpal IV is the longest, followed by metacarpals III and II. Metacarpal V of the right hand is a small dumbbellshaped bone. The conical ungual on digit I is well developed, and digit I is set at an oblique angle to the main axis of the manus. The ungual phalanx is quite characteristic of basal iguanodontoids: it is straight, triangular, while in more derived iguanodontians the rst phalanx of digit I may be lacking (Norman 2004). Ungual phalanx of digit II is large and subtriangular in the dorsal view; unlike the holotype of Bolong (Wu and Godefroit 2012), the ungual II is smaller than ungual I. The left digit V has two phalanges. Other phalanges are poorly preserved. The morphology of the manus is similar to the holotype of Bolong (Wu and Godefroit 2012), the rst digit has an enlarged, divergent, spine-like ungual, the middle three digits form a rather compact unit and the fth digit is apparently long, exible and opposable. This is similar to the condition in Iguanodon (Norman 1986, 2004), and different from that of many basal iguanodontians, such as Tenontosaurus (Forster 1990b) and Zalmoxes (Weishampel et al. 2003).

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

Pelvic girdle and hindlimbs Both ilia are preserved, but obscured by overlaying remains. The anterior portions of preacetabular process of both ilia are exposed in the medial view (Figures 2 and 13). The process directs and tapers anteriorly; few anatomical details can be ascertained from the material. Both pubes are preserved in the medial view (Figures 2 and 13). The prepubic process of the pubis is slender, concave along its dorsal margin and expanded at its distal end. It is remarkably similar to those of Iguanodon bernissartensis (Norman 1980) and Altirhinus (Norman 1998). The pubic shaft is short and tapering, it is probably terminated about mid-way along the shaft of the ischium. This is a derived character for a non-hadrosaurid iguanodontian (Norman 2004). Both ischia are preserved in the medial view (Figures 2 and 13). The acetabular margin is visible on the right ischium and is gently concave in the medial view. In the medial view, the ischial shaft is straight to slightly arched dorsally and becomes deeper along its length. The condition of the distal end is difcult to ascertain; it looks like the distal end is expanded ventrally. The obturator process is well developed on the ventral margin of the shaft. Both femora are present (Figures 2 and 14). The femur is robust and straight. The fourth trochanter is pendant and situated at approximately midlength of the shaft. Beneath the fourth trochanter, the shaft is curved posteriorly. The distal end of the femur is slightly expanded anteroposteriorly with respect to the shaft. The tibia is straight and slightly shorter than the femur (Figure 14, Table 1). The proximal end of the tibia is broad and expanded. The distal end is transversely expanded to

248 W. Zheng et al. form the inner and outer malleoli. The outer malleolus is separated from the inner by a step and projects more distally. The distal bula is articulated against the anterior surface of the outer malleolus. The bula is a straight, elongated and rod-like bone attached to the lateral surface of the tibia. Its proximal end is anteroposteriorly expanded. The shaft tapers towards its distal end. Metatarsals II IV are tightly bound together into a single broad metapodial unit (Figure 15). Metatarsal II is laterally compressed and anteroposteriorly expanded. Its mid shaft is slender, and the distal end is also anteroposteriorly expanded. Metatarsal III is the longest of the three elements, and straight. The proximal end of metatarsal IV is expanded. About half-way down the shaft, a vertical ridge is bound by ligaments to metatarsal III. Only three phalanges of the right foot are preserved. They are broad and have expanded proximal and distal ends. The metatarsals are more slender than that in the holotype of Bolong. The metatarsals are stout in YZH-001 with a ratio of metatarsal III length to femur equal to 0.18; however, the metatarsals are distorted (Wu and Godefroit 2012). embryonic specimen of Hypacrosaurus (Gilmore 1924; Horner and Currie 1994). The jugal is more robust in the adult Bolong. The overall jugal robustness is also reported variable rquez 2011). ontogenetically in hadrosauroids (Prieto-Ma The morphology of scapula in M8812 is very similar to that in YZH-001, whereas the scapula is more slender in M8812, indicating that the scapula becomes robust with age. Similarly, the scapula of basal iguanodontian Tenontosaurus is broader in adults (Forster 1990b). The radius and ulna are more robust in the adult Bolong, whereas the robustness of tibia remains almost the same. The proportions of fore- and hindlimb also change. Although the humerus and femur are not complete in the holotype of Bolong, the forelimb appears proportionally very short and more closely resembles the condition encountered in Mantellisaurus athereldensis (humerus is 58 56% of femoral length). In ZMNH M8812, the humerus is approximately 81% of femoral length. The hindlimb forelimb ratio is approximately 1.46. The length of hindlimb is the sum of the lengths of the femur, tibia and metatarsal III; and the length of forelimb is the sum of the lengths of the humerus, radius and metacarpal III as in Norman (1980). The ratio is close to Iguanodon bernissartensis, more like a quadrupedal dinosaur (Norman 1980). The index of forelimb proportions, which is multiplied the ratios of the lengths of radius/humerus and metacarpal III/humerus, is approximately 0.21. This ratio is very close to that in Dollodon, which also suggests that the M8812 is also closer to that of typical quadruped (Norman 1980).

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

Discussion Ontogenetic changes The description of this juvenile specimen allows the study of ontogenetic of Bolong for the rst time. The body length of this individual is , 50 cm. Osteohistological study suggests a very early developmental stage of this individual. In the holotype of Bolong, the caudal ribs are not fused to the centra, indicating that sexual maturity was probably not completely achieved. However, the complete closure of the sutures between the neural arches and centra throughout the vertebral column is suggestive of skeletal maturity. The body length was estimated approximately 3 m for the holotype of Bolong (Wu and Godefroit 2012). The body size is relatively small than Jinzhousaurus, which has complete closure of the sutures between the neural arches and centra throughout the vertebral column, the body length is approximately 5 5.5 m (Wang et al. 2010). The number of tooth positions in the maxilla and dentary increases during ontogeny as in other iguanodon bner tians (Weishampel et al. 2003; Horner et al. 2004; Hu and Rauhut 2010); the morphology of teeth is very similar in juvenile and adult Bolong, as reported in basal ornithopod Talenkauen (Egerton et al. 2013) and the hadrosaurid Hypacrosaurus (Evans 2010, g. 12; Horner and Currie 1994, g. 21.23). The teeth in juvenile hadrosaurids are reported narrower than adult individuals, although the height-to-width ratio of tooth crown remains almost the same in juveniles and adults of Bactrosaurus rquez 2011), the dental teeth are more (Prieto-Ma anteroposterorly narrower in adults than those in an

Phylogenetic analysis The phylogenetic placement of this specimen within iguanodontians was evaluated quantitatively by adding it to the matrix of McDonald (2012), with the holotype of Bolong (YHZ-001) (Wu et al. 2010) and Proa (McDonald et al. 2012) also included. The character scores for YHZ-001 are based on the detailed description by Wu and Godefroit (2012). The matrix we used here consisted of 69 operational taxonomic units (OTUs) and 135 characters. The analysis was carried out using traditional search with the tree bisection reconnection algorithm in TNT version 1.1 (Goloboff et al. 2008), with all characters equally weighted and 12 characters (10, 14, 20, 25, 46, 67, 81, 82, 83, 100, 127, 130) ordered. Starting trees were Wagner trees with a random seed of 1; 9999 replicates were used with 10 trees saved per replication. Five OTUs (Camptosaurus valdensis, Draconyx, NHMUK R8676, Delapparentia and Glishades) were excluded (McDonald 2012; McDonald et al. 2012). The analysis resulted in 10,750 most parsimonious trees (MPTs) of 415 steps. The strict consensus tree was very poorly resolved, with nearly the whole of Iguanodontia in an

Historical Biology 249

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

Figure 16. The 50% majority rule consensus tree of 10,750 MPTs recovered from a traditional (heuristic) search in TNT. Figures below nodes represent the percentage of MPTs in which the node was recovered.

unresolved polytomy. The 50% majority rule consensus tree of 10,750 MPTs (415 steps) was recovered from a traditional search (Figure 16). Both ZMNH M8812 and YHZ-001 are recovered as basal styracosternans, within a node that is very poorly resolved. Bolong is basal to most other iguanodontians from east-central Asia (such as Altirhinus, Equijubus, Probactrosaurus or Jinzhousaurus), similar to the result in Wu and Godefroit (2012). The phylogenetic characters used by McDonald (2012) for inferring iguanodontian relation-

ships are not affected by ontogeny in Bolong, as in Bactrosaurus that the juvenile and subadult specimens can provide a substantial amount of reliable information for rquez 2011). phylogenetic inference (Prieto-Ma Conclusions A juvenile specimen of Bolong yixianensis was described, which has signicantly increased the anatomical knowl-

250 W. Zheng et al. edge of this basal iguanodontian ornithopod. The specimen revealed one additional autapomorphy for Bolong yixianensis. The lingual face of the crown is bounded by thickened mesial and distal margins and is bisected by a prominent median principal ridge. Osteohistologic studies of the femur indicate that this individual was in the nestling stage when it died, probably only several months after it hatched. Thus, ZMNH M8812 is the rst early juvenile iguanodontian specimen reported from east Asia, as such, an important specimen for future ontogenetic studies of basal iguanodontians. The comparison between the juvenile and holotype of Bolong yixianensis revealed the following ontogenetic variations in this taxon: the adult maxilla is deep and the apex is more distally positioned, the adult jugal is more robust, the tooth rows of both maxilla and dentary are less in the juvenile individual, the scapula is more robust in the adult, the radius and ulna are more robust in the adult and became proportionally shorter relative to the hindlimb; the adult metatarsals are proportionally shorter than the juvenile. The following characters, previously used in the diagnosis of Bolong yixianensis, are variable ontogenetically: the primary ridge is deected distally on maxillary crowns, ulna and radius are proportionally short and robust, metatarsals are proportionally short. Furthermore, this study suggests that, in Bolong yixianensis, the phylogenetic characters used by McDonald (2012) for inferring iguanodontian relationships are not affected by ontogeny. Thus, juvenile specimens may provide reliable information in phylogenetic analysis. Acknowledgements
We are grateful to David Varricchio and Frankie Jackson (Montana State University, USA) for reading the manuscript and offering many valuable suggestions. We thank Yuqing Zhang and Chaohe Yu for preparing the specimen and Dr Junchang Lu (Institute of Geology, Chinese Academy of Geological Sciences, Beijing) for helping in the process of preparation. The thin sections of the femur were made at Museum of the Rockies, Montana. We thank Anusuya Chinsamy-Turan (University of Cape Town, South Africa) for helpful comments in our osteohistology study.
Dollo L. 1888. Iguanodontidae et Camptonotidae. CR Acad Sci. 106:775777. Egerton VM, Novas FE, Dodson P, Lacovara K. 2013. The rst record of a neonatal ornithopod dinosaur from Gondwana. Gondwana Res. 23(1):268271. Evans DC. 2007. Ontogeny and evolution of Lambeosaurine dinosaurs (Ornithischia: Hadrosauridae). Unpublished Ph.D. Dissertation: University of Toronto. Evans DC. 2010. Cranial anatomy and systematics of Hypacrosaurus altispinus, and a comparative analysis of skull growth in lambeosaurine hadrosaurids (Dinosauria: Ornithischia). Zool J Linn Soc. 159(2):398434. Forster CA. 1990a. Evidence for juvenile groups in the ornithopod dinosaur Tenontosaurus tilletti Ostrom. J Paleontol. 64(1):164 165. Forster CA. 1990b. The postcranial skeleton of the ornithopod dinosaur Tenontosaurus tilletti. J Vert Paleontol. 10(3):273294. Galton PM. 1974. The ornithischian dinosaur Hypsilophodon from the Wealden of the Isle of Wight. Bull Br Mus Nat Hist. 25(1):1151. Gilmore CW. 1924. On the skull and skeleton of Hypacrosaurus, a helmet-crested dinosaur from the Edmonton Cretaceous of Alberta. Can Depart Mines Geol Surv Can Bull. 38:4964. Gilmore CW. 1933. On the dinosaurian fauna of the Iren Dabasu Formation. Bull Am Mus Nat Hist. 67:2378. Godefroit P, Dong Z-M, Bultynck P, Li H, Feng L. 1998. New Bactrosaurus (Dinosauria: Hadrosauroidea) material from Iren Dabasu (Inner Mongolia, P.R. China). Bull Inst R Sci Nat Belg Sci Terra. 68:3 70. Goloboff PA, Farris JS, Nixon KC. 2008. TNT, a free program for phylogenetic analysis. Cladistics. 24(5):774786. bner TR, Rauhut OWM. 2010. A juvenile skull of Dysalotosaurus Hu lettowvorbecki (Ornithischia: Iguanodontia), and implications for cranial ontogeny, phylogeny, and taxonomy in ornithopod dinosaurs. Zool J Linn Soc. 160(2):366396. Horner JR, Currie PJ. 1994. Embryonic and neonatal morphology and ontogeny of a new species of Hypacrosaurus (Ornithischia, Lambeosauridae) from Montana and Alberta. In: Dinosaur eggs and babies. Cambridge: Cambridge University Press. RERO DOC [http://doc.rero.ch/oai2d.py/] (Switzerland) ER. p. 312 336. ` s A, Padian K. 2000. Long bone histology of the Horner JR, De Ricqle hadrosaurid dinosaur Maiasaura peeblesorum: growth dynamics and physiology based on an ontogenetic series of skeletal elements. J Vert Paleontol. 20(1):115129. Horner JR, Weishampel DB, Forster CA. 2004. Hadrosauridae. In: The Dinosauria. 2nd ed. Berkeley, CA: University of California Press. p. 438 463. Irmis RB. 2007. Axial skeleton ontogeny in the Parasuchia (Archosauria: Pseudosuchia) and its implications for ontogenetic determination in archosaurs. J Vert Paleontol. 27(2):350361. Liu Y, Liu Y, Ji Sa, Yang Z. 2006. U-Pb zircon age for the Daohugou Biota at Ningcheng of Inner Mongolia and comments on related issues. Chin Sci Bull. 51(21):26342644. Marsh OC. 1881. Principal characters of American Jurassic dinosaurs. Pt V. Am J Sci (Series 3). 21:417 423. McDonald AT. 2012. Phylogeny of basal iguanodonts (Dinosauria: Ornithischia): an update. PLoS ONE. 7(5):e36745. lez E, Mampel L, Kirkland JI, Alcala L. 2012. An McDonald AT, Esp unusual new basal iguanodont (Dinosauria: Ornithopoda) from the Lower Cretaceous of Teruel, Spain. Zootaxa. 3595:6176. Norman DB. 1980. On the ornithischian dinosaur Iguanodon bernissar m Inst R Sci Nat Belg. 178:1103. tensis from Belgium. Me Norman DB. 1986. On the anatomy of Iguanodon athereldensis (Ornithischia: Ornithopoda). Bull Inst R Sci Nat Belg. 56:281372. Norman DB. 1996. On Mongolian ornithopods (Dinosauria: Ornithischia). 1. Iguanodon orientalis Rozhdestvensky 1952. Zool J Linn Soc. 116(3):303315. Norman DB. 1998. On Asian ornithopods (Dinosauria: Ornithischia). 3. A new species of iguanodontid dinosaur. . Zool J Linn Soc. 122(1 2):291348. Norman DB. 2002. On Asian ornithopods (Dinosauria: Ornithischia). 4. Probactrosaurus Rozhdestvensky, 1966. Zool J Linn Soc. 136(1):113 144. Norman DB. 2004. Basal Iguanodontia. In: The dinosauria. 2nd ed. Berkeley, CA: University of California Press. p. 413437.

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

References
Barrett PM, Butler RJ, Wang X-L, Xu X. 2009. Cranial anatomy of the iguanodontoid ornithopod Jinzhousaurus yangi from the Lower Cretaceous Yixian Formation of China. Acta Palaeontol Pol. 54(1):3548. Brink KS, Zelenitsky DK, Evans DC, Therrien F, Horner JR. 2010. A subadult skull of Hypacrosaurus stebingeri (Ornithischia: Lambeosaurinae): anatomy and comparison. Hist Biol. 23(1):63 72. Brochu CA. 1996. Closure of neurocentral sutures during crocodilian ontogeny: implications for maturity assessment in fossil archosaurs. J Vert Paleontol. 16(1):49 62. Cerda IA, Chinsamy A. 2012. Biological implications of the bone microstructure of the Late Cretaceous Ornithopod Dinosaur Gasparinisaura cincosaltensis. J Vert Paleontol. 32(2):355368.

Historical Biology 251

Owen R. 1842. Report on british fossil reptiles. Part II. Rep Br Assoc Adv Sci. 11:60204. rquez A. 2011. Cranial and appendicular ontogeny of Prieto-Ma Bactrosaurus johnsoni, a hadrosauroid dinosaur from the Late Cretaceous of northern China. Palaeontology. 54(4):773792. Scheetz RD. 1999. Osteology of Orodromeus makelai and the phylogeny of basal ornithopod dinosaurs. Bozeman, MT: Montana State University. Seeley HG. 1887. On the classication of the fossil animals commonly named Dinosauria. Proc R Soc Lond. 43:165171. Shibata M, Jintasakul P, Azuma Y. 2011. A new Iguanodontian dinosaur from the Lower Cretaceous Khok Kruat Formation, Nakhon Ratchasima in Northeastern Thailand. Acta Geol Sin Engl Ed. 85(5):969976. Wang X, Pan R, Butler RJ, Barrett PM. 2010. The postcranial skeleton of the iguanodontian ornithopod Jinzhousaurus yangi from the Lower Cretaceous Yixian Formation of western Liaoning, China. Earth Environ Sci Trans R Soc Edinb. 101(02):135 159. Wang X-L, Wang Y-Q, Zhang F-C, Zhang J-Y, Zhou Z-H, Jin F, Hu Y-M, Gu G, Zhang H-C. 2000. Vertebrate biostratigraphy of the Lower Cretaceous Yixian Formation in Lingyuan, western Liaoning and its neighboring southern Nei Mongol (Inner Mongolia), China. Vert PalAsiatic. 38(2):8199. Wang X, Xu X. 2001. A new iguanodontid (Jinzhousaurus yangi gen. et sp. nov.) from the Yixian Formation of western Liaoning, China. Chin Sci Bull. 46(19):1669 1672. Wang X, Zhou Z, He H, Jin F, Wang Y, Zhang J, Wang Y, Xu X, Zhang F. 2005. Stratigraphy and age of the Daohugou Bed in Ningcheng, Inner Mongolia. Chin Sci Bull. 50(20):2369 2376. Weishampel DB, Barrett PM, Coria RA, Le Loeuff J, Xu X, Zhao X, Sahni A, Gomani EMP, Noto CR. 2004. Dinosaur distribution. In: The dinosauria. 2nd ed. Berkeley, CA: University of California Press. p. 517606. Weishampel DB, Jianu C-M, Csiki Z, Norman DB. 2003. Osteology and phylogeny of Zalmoxes (n. g.), an unusual Euornithopod dinosaur from the latest Cretaceous of Romania. J Syst Palaeontol. 1(2):65123. Werning S. 2012. The ontogenetic osteohistology of Tenontosaurus tilletti. PLoS One. 7(3):e33539. Wu W-H, Godefroit P, Hu D-Y. 2010. Bolong yixianensisgen gen. et sp. nov.: a new iguanodontoid dinosaur from the Yixian Formation of western Liaoning, China. Geol Resour. 19(2):127133. Wu W, Godefroit P. 2012. Anatomy and relationships of Bolong yixianensis, an Early Cretaceous Iguanodontoid dinosaur from Western Liaoning, China. In: Bernissart dinosaurs and Early Cretaceous Terrestrial ecosystems. Bloomington: Indiana University Press. p. 292333. You H-L, Luo Z-X, Shubin NH, Witmer LM, Tang Z-l, Tang F. 2003. The earliest-known duck-billed dinosaur from deposits of late Early Cretaceous age in northwest China and hadrosaur evolution. Cret Res. 24(3):347355. You H, Ji Q, Li D. 2005. Lanzhousaurus magnidens gen. et sp. nov. from Gansu Province, China: the largest-toothed herbivorous dinosaur in the world. Geol Bull China. 24(9):785794.

Downloaded by [Harbin Institute of Technology] at 10:35 01 April 2014

Appendix: The character scoring for ZMNH M8812 and YHZ-001 in the matrix by McDonald (2012)
ZMNH M8812: ????? ????1 ?0?01 1?1?? 0???? ????1 ????? ????? 00?21 ????? ????0 ?1??? ????? ????? ????? ????? 10100 11210 100?? ????1 10000 111?? ????? 11111 ?0?01 ????? ?????. YHZ-001: ??1?1 11?11 ?0?03 1?1?0 00?1? ???11 101?? 1111? 10??1 22030 11??0 ????? ??1?1 ????? ????? ????? 11100 11210 10001 0???1 10000 1111? ???10 ????2 10??? ?1??? 2????.