A new hadrosauroid dinosaur from the Late Cretaceous of Tianzhen, Shanxi Province, China

Xu Shi-ChaoYOu Hai-LuWAnG Jia-WeiWAnG Suo-ZhuYI JianJIA Lei,2

(1Shanxi Museum of GeologyTaiyuan 030024, China)

(2Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences,Institute of Vertebrate Paleontology and Paleoanthropology,Chinese Academy of SciencesBeijing 100044, China * Corresponding author: youhailu@ivpp.ac.cn)

(3Faculty of Geology,Lomonosov Moscow State University Moscow 119991, Russian Federation)

A new hadrosauroid dinosaur from the Late Cretaceous of Tianzhen, Shanxi Province, China

Xu Shi-Chao1YOu Hai-Lu2*WAnG Jia-Wei3WAnG Suo-Zhu1YI Jian1JIA Lei1,2

(1Shanxi Museum of GeologyTaiyuan 030024, China)

(2Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences,Institute of Vertebrate Paleontology and Paleoanthropology,Chinese Academy of SciencesBeijing 100044, China * Corresponding author: youhailu@ivpp.ac.cn)

(3Faculty of Geology,Lomonosov Moscow State University Moscow 119991, Russian Federation)

A new non-hadrosaurid hadrosauroid dinosaur (Datonglong tianzhenensisgen. et sp. nov.) is reported. The new taxon is recovered from the Upper Cretaceous Huiquanpu Formation of Tianzhen County, Shanxi Province in northern China, and represented by an almost complete right dentary with dentition. Different from all other hadrosauroids,Datonglongpossesses two functional teeth in each alveolus, and the pattern of ridge development on the lingual surface of its dentary crown shows a unique combination of character states (for example: distally offset primary ridge; well-developed secondary ridge; no additional ridge(s); slightly distally curved apical half of primary ridge). Comparative studies indicate advanced non-hadrosaurid hadrosauroids experienced a complex pattern in the evolution of their dentary, especially dentary dentition. Derived hadrosaurid features occurred frequently in these taxa, such as high height/width ratio of tooth crown inBactrosaurus, one primary and one faint ridges inGilmoreosaurus, median placed primary ridge inZhanghenglong, rostrally inclined coronoid process inNanningosaurus, and two functional teeth in each alveolus inDatonglong. This implies incredible diversities and attempts close to the origin of Hadrosauridae and dif fi culties to elucidate their phylogenetic relationships.

Tianzhen, Shanxi; Late Cretaceous; dinosaur, hadrosauroid

1 Introduction

The Huiquanpu Formation was established by the Regional Geological Survey Team ofHebei Province in 1959. It refers to the over 200 meter thick terrestrial deposits bounded by the pre-Cambrian rocks and the Miocene basalts, and is distributed in an area of about 150 km2around the boundary of Tianzhen County of Shanxi Province and Yangyuan County of Hebei Province in northern China, with an age assignment of Eocene. Liu (1983) and Chen et al. (1983) reassigned its age to Late Cretaceous based on discoveries of dinosaurs, especially Hadrosauridae dinosaurs. In 1989 and 1991-1994, Shijiazhuang Economic University of Hebei Province excavated over 2300 dinosaur specimens mainly in the Kangdailiang Quarry of Tianzhen County, Shanxi Province (Pang et al., 1996), and two new dinosaurs have been established, namely an ankylosaurTianzhenosaurusyoungi Pang & Cheng, 1998, and a sauropodHuabeisaurus allocotusPang & Cheng, 2000. In 1998, another ankylosaurShanxia tianzhenensisBarrett et al., 1998 was also reported from the Wujiashan Quarry of Tianzhen County, about 7 km northeast of the previous Kangdailiang Quarry. These two ankylosaurs may be distinct from each other (Upchurch and Barrett, 2000; Thompson et al., 2012), or represent the same taxon (Sullivan, 2000); whileTianzhenosaurusmay be synonymous withSaichania(Sullivan, 1999). Recent restudy ofHuabeisaurus allocotusshows this taxon is probably a member of the Cretaceous East Asian endemic clade of Euhelopodidae (D’Emic et al., 2013).

Pang et al. (1996) and Pang and Chen (2000, 2001) also mentioned the existence of theropod cf.Szechuanosaurus campiYoung, 1942 and hadrosaurid cf.Shantungosaurussp. in the Huiquanpu Formation. The former has been regarded as a nomen dubium (Carrano et al., 2012); while the assignment of fragmentary material to cf.Shantungosaurussp. is based on observation of their tibia similarity (Pang and Cheng, 2000, 2001).

Fig. 1 Locality ofDatonglong tianzhenensisgen. et sp. nov. (SXMG V 00005) A. Shanxi Province in China; B. Tianzhen County in Shanxi Province

Here we report a new hadrosauroid specimen recovered from the Kangdailiang Quarry by the Shanxi Museum of Geological and Mineral Science and Technology (now Shanxi Museum of Geology, SXMG) in 2008 (Fig. 1). Although the specimen is represented only bya right dentary with dentition, it provides numerous important anatomical features to depict its taxonomic status and systematic relationship. Based on our anatomical observation and taxonomic comparison, the new specimen is distinct from all previous known hadrosauroid dinosaurs, represents a new taxon (Datonglong tianzhenensisgen. et sp. nov.), and belongs to an advanced non-hadrosaurid hadrosauroid.

Institutional abbreviations AMNH, American Museum of Natural History (New York City, New York); LPM, Liaoning Paleontological Museum (Beipiao, Liaoning); NHMG, Natural History Museum of Guangxi Zhuang Autonomous Region (Nanning, Guangxi); SBDE, Sino-Belgium Dinosaur Expedition, Inner Mongolia Museum (Hohhot, Nei Mongol Autonomous Region); SXMG, Shanxi Museum of Geology (Taiyuan, Shanxi); XMDFEC, Xixia Museum of Dinosaur Fossil Eggs of China (Xixia, Henan).

2 Systematic paleontology

Dinosauria Owen, 1842

Ornithischia Seeley, 1887

Ornithopoda Marsh, 1881

Iguanodontia Dollo, 1888 sensu Sereno, 2005

Ankylopollexia Sereno, 1986 sensu Sereno, 2005

Styracosterna Sereno, 1986 sensu Sereno, 2005

Hadrosauriformes Sereno, 1997 sensu Sereno, 1998

Hadrosauroidea Sereno, 1986 sensu Sereno, 2005

Datonglonggen. nov.

Datonglong tianzhenensissp. nov.

Holotype SXMG V 00005, almost complete right dentary with dentition.

Etymology The generic name “Datong” is after “Datong City”, and Tianzhen is a county belonging to it; “l(fā)ong” means “dragon” in Chinese. The speci fi c name “Tianzhen” is after “Tianzhen County”, where the current material is discovered.

Locality and horizon Kangdailiang Quarry, Tianzhen County, Datong City, Shanxi Province, China. Huiquanpu Formation, Upper Cretaceous.

Differential diagnosis (for genus and species by monotypy) Advanced non-hadrosaurid hadrosauroid with one autapomorphy (two functional teeth in each alveolus along at least the mid- and caudal dentary occlusal plane) and the following unique combination of character states in the pattern of ridge development on the enameled lingual surface of dentary tooth crown: 1) distally offset primary ridge, 2) well-developed secondary ridge, 3) no additional ridge(s), 4) slightly distally curved apical half of primary ridge.

Description An almost complete 34 cm long right dentary is preserved, missing its rostral end including the rostral end of the tooth row (Fig. 2). The preserved ramus of thedentary is rectangular in lateral view, with roughly parallel dorsal and ventral margins, 14 cm high at the mid portion with the occlusal plane or 10.5 cm high without it (Fig. 2A). The tooth row (28 cm long) ends caudally right medial to the caudal margin of the coronoid process (Fig. 2B). The 8 cm high coronoid process is almost perpendicular and slightly caudodorsallydirected to the ramus, and the dentary height along it is 17 cm. Although the tip of the coronoid process is not preserved, its dorsal portion is expanded at least rostrally as evidenced by its broken contour, and its craniocaudal width at base is 4.8 cm. Its lateral surface bulges slightly, while the medial one is flat, bearing a faint vertical ridge limiting its caudal two fi fths with further ridges on it. In dorsal view, the caudal half of the ramus curves laterally, with caudally increasing distance between the tooth row and the lateral surface of the ramus, ending in an about 3 cm wide horizontal shelf separating the coronoid process and the caudal end of the tooth row (Fig. 2C). In caudal view, a triangular fossa is bounded by the coronoid process (Fig. 2D). The fossa extends ventrally to the mandibular adductor fossa. Rostral to the adductor fossa, the Meckelian groove is deep and long, and progressively decreases its dimensions toward the preserved rostral end.

Fig. 2 Photos of right dentary ofDatonglong tianzhenensisgen. et sp. nov. (SXMG V 00005) A. lateral view; B: medial view. C: dorsal view; D. caudal view; E. close-up of partial dentition in B; E is not in scale

The tooth row is largely preserved, with 17 alveoli full of teeth plus five successive most caudal alveoli bearing several scattered teeth. At least another fi ve alveoli exist rostral to the above 22 alveoli judging by the preserved length and the existence of two broken teeth in this portion. Therefore, a total of at least 27 alveoli are expected for this animal; and theactual number is probably less than 30 based on the general alveolar contour, which reduces its depth rapidly rostrodorsally. In medial view, a row of neurovascular foramina slightly extends caudodorsally along the mid-caudal portion of the ramus, and in this portion about three quarters of the dorsoventral depth of the dentary ramus is occupied by the alveoli. The partial septum dividing the fourth and fi fth most caudal alveoli is exposed, and it is straight and relatively thick (2 mm).

The teeth are best exposed in the mid-caudal portion of the tooth row. In medial view, two or three teeth are vertically aligned in each alveolus (Fig. 2B). In lateral view, each alveolus bears two functional teeth, of which the upper one is large and obviously the same as the uppermost one seen medially, while the lower one should be the remaining worn facet of the earlier functional tooth (Fig. 2A). Therefore, three or four teeth are held in each alveolus. Both worn surfaces in each alveolus are slightly concave, and in some alveoli the upper one is not continuous with the lower one, with the latter placed more medially.

The teeth are large, and their morphology can be best shown in the tenth functional and eleventh replacement positions counting from the caudal (Fig. 2E). A distally offset primary ridge exists on the enameled lingual side in teeth, and is accompanied by a welldeveloped secondary ridge in the mesial half of the crown. These two roughly parallel ridges divide the lingual crown surface into three zones, without evidence of any subsidiary ridges on it. The ventral halves of these two ridges are vertical, while the dorsal halves curve slightly caudodorsally. The unerupted tooth is diamond-shaped, with the height of 5.5 cm and maximum width of 2.0 cm, a ratio of 2.75. One scattered tooth attached to the third last alveolus exposes its mesial margin, where small mammillated papillae occur apically.

Comparison In the evolution from non-hadrosaurid hadrosauroids to hadrosaurids, their dentaries acquired several key innovations. In all hadrosaurids, the tooth row ends caudally to the caudal margin of the coronoid process (Xing et al., 2014: character 56) and the coronoid process inclines rostrally (Xing et al., 2014: character 48). In addition, the dentary dental battery bears more than 30 alveoli (Xing et al., 2014: character 1), and the unworn crown in the mid-tooth row is elongate lanceolate with a height/width ratio greater than 3.1 (Sues and Averianov, 2009). All these features are not present inDatonglong. Therefore,Datonglongdoes not belong to Hadrosauridae, and our comparison will focus on nonhadrosaurid hadrosauroids, especially those from the Late Cretaceous of China. Based on Wang et al. (2013), Xing et al. (2014), and Tsogtbaatar et al. (2014), 15 Late Cretaceous nonhadrosaurid hadrosauroid genera have been reported, with eight from China (TaniusWiman, 1929;BactrosaurusGilmore, 1933;GilmoreosaurusBrett-Surman, 1979;NanyangosaurusXu et al., 2000;ShuangmiaosaurusYou et al., 2003;NanningosaurusMo et al., 2007;YunganglongWang et al., 2013;ZhanghenglongXing et al., 2014), three from North America (ClaosaurusMarsh, 1890 ;EolambiaKirkland, 1998;ProtohadrosHead, 1998), two from Europe (TelmatosaurusNopcsa, 1903;TethyshadrosDalla Vecchia, 2009), one from Central Asia (LevnesoviaSues & Averianov, 2009), and one from Mongolia (PlesiohadrosTsogtbaatar et al., 2014). Among these eight Chinese taxa, three (Tanius,Nanyangosaurus, andYunganglong) do not preserve comparable parts withDatonglong.

Nanningosaurus, recovered from the Upper Cretaceous red beds of Guangxi in southern China, was originally reported as a lambeosaurine hadrosaurid (Mo et al., 2007). Recently, it was recovered as a non-hadrosaurid hadrosauroid (Xing et al., 2014). A left dentary and an isolated dentary tooth are included in the holotype (NHMG 8142). It has a high coronoid process that inclines slightly rostrally and ends slightly caudal to the coronoid process, definitely derived features typical for hadrosaurids and different from the conditions inDatonglong. However, two ridges are present on the lingual crown of dentary tooth as inDatonglong. The primary ridge is sinuous as seen inTsintaosaurus.

An isolated left dentary (LPM 0166) is referred toShuangmiaosaurus gilmorei(You et al., 2003a). InShuangmiaosaurus, the coronoid process inclines caudodorsally and the tooth row ends in a position medial to the middle of the coronoid process. Moreover, the dorsal edge of the dentary ramus slightly bows dorsally and the 27 alveoli clearly incline caudodorsally. All these features are not present in similar-sizedDatonglong.

Bactrosauruswas originally studied by Gilmore (1933) based on material from a bonebed in Quarry 141 (Johnson’s Quarry) collected by Central Asiatic Expeditions of the American Museum of Natural History. AMNH 6553 is designated as the holotype, and it contains more than one individual including left and right dentaries. Based on Gilmore (1933), 12 dentaries have been collected in this bonebed, and more than half of them pertain to juveniles. However, Gilmore’s description of dentary seems to be based on AMNH 6353 (a right dentary), with a drawing of its medial and dorsal views. The length of adult dentary is 222 mm, and the length of the longest tooth row is 152 mm with 23 alveoli. Each alveolus holds one replacement tooth, and at most two functional teeth. A large unworn tooth crown has a ratio of 3.3 (33/10 mm).

Prieto-Marquez (2011) studied the ontogeny ofBactrosaurus. He notices that in the adult dentary (AMNH 6553: 23 cm long) there are as many as three teeth per alveolus arranged dorsoventrally at mid-length of the dental battery with two functional teeth. The tooth crowns have a height/width ratio of 3.1-3.2 as in juvenile specimens; in contrast to 2.75 inDatonglong(55/20 mm). The position of the primary ridge is slightly or modestly offset from the midline.

Godefroit et al. (1998) studiedBactrosaurusfrom another bonebed excavated by SBDE in 1995, within 1 km to AMNH Quarry 141. It contains several hundred bones of at least four hadrosaur individuals, including a left dentary (SBDE 95E5/12). Godefroit also demonstrated that all hadrosauroids in Quarry 141 belong toBactrosaurusas Gilmore originally suggested. SBDE 95E5/12 is about the same size as the holotype dentary (AMNH 6553), and bears 20 alveoli. In contrast, around 30 alveoli have been estimated forDatonglong; however, the length of dentary inDatonglongis about 1.5 times as that in adultBactrosaurus. Godefroit et al. (1998) described that each alveolus is composed of one functional tooth and at least tworeplacement teeth. However, based on the illustration, two or three teeth can be seen medially, and two functional teeth are visible laterally in the preserved mid- and caudal portion of the tooth row. Here, based on our observation inDatonglong, the lower functional tooth should be from the earlier worn facet, but not from any tooth can be seen medially. Therefore, in bothBactrosaurusandDatonglong, at most two replacement and two functional teeth are present in one alveolus. However, two functional teeth seem to persist along the entire tooth tow inDatonglong, but not in the rostral portion of the tooth row inBactrosaurusas shown in AMNH 6553. As inDatonglong,Bactrosaurusalso has two ridges, one primary and one relatively well developed secondary ridge on the lingual tooth crown; however inBactrosaurusthere are evidence of other faint ridges in the adults, and a third caudal ridge appears in caudal most teeth. The primary ridges may curve slightly caudally inBactrosaurusas inDatonglong.

Therefore, the crown ratio, the number of alveoli, and the condition of functional teeth in rostral portion of tooth row are different betweenBactrosaurusandDatonglong. The details of ridge development pattern in the dentary tooth crown are also different.

Prieto-Marquez and Norell (2010) redescribed the second hadrosauroidGilmoreosaurus mongoliensisfrom the Iren Dabasu Formation in Irenhot. This species is represented by at least four individuals from quarries 145 and 149. Among the material, only a partial right dentary (AMNH FARB 30654) and an isolated dentary tooth (AMNH FARB 30661) are preserved. The dentary mainly consists of the rostral and middle portion without the coronoid process and dentition (295 mm long and 71 mm high), and it is hard to tell de fi nite diagnostic differences between it andDatonglong, although the latter seems to be more robust. In contrast, the worn half crown of an isolated dentary tooth possesses only one ridge that lies off center, different from the condition inDatonglong, which has two ridges.Gilmoreosaurusprobably possesses one primary and one faint ridges on its dentary tooth crown (personal communication with Xing Hai).

Zhanghenglongis a recently described hadrosauroid from the Santonian Majiacun Formation of Xixia Basin, Henan Province in central China (Xing et al., 2014). Its holotype (XMDFEC V0013) preserves an incomplete, disarticulated cranium, including a right dentary with dentition. InZhanghenglong, the tooth row bows dorsally and its long axis is nearly parallel to the lateral side of the dentary ramus, unlike the horizontal and diverged condition inDatonglong. The 26 alveoli bear one functional tooth at the rostral and caudal portions, two in the midsection, and even three in the 17thtooth alveolus; while inDatonglong, two functional teeth seem to persist along the entire tooth row. InZhanghenglong, the primary ridge is situated median (about 40%) or distally offset on the lingual crown of the dentary, a clear secondary ridge is on the mesial half, and additional faint ridge(s) may appear close to the mesial edge; in contrast inDatonglong, all primary ridges are placed in the distal halves, and no additional ridge(s) are evident besides the secondary one. The dentary tooth crown has an approximate height/width ratio of 2.36 inZhanghenglong, and less than 2.75 as inDatonglong.

The latest reported non-hadrosaurid hadrosauroid isPlesiohadros djadokhtaensisfrom the Campanian of Mongolia (Tsogtbaatar et al., 2014).Plesiohadrospreserves a left dentary with dentition. The dentary is relatively slender with a straight ramus in occlusal view, while that ofDatonglongis relatively robust with a laterally curved ramus. Besides the primary and mesial secondary ridges, a distal secondary ridge occurs in some teeth inPlesiohadros, while only two ridges exist inDatonglong.

3 Discussion

Several dentition features show thatDatonglongis an advanced taxon among nonhadrosaurid hadrosauroids. The dentary teeth became miniaturized in the evolution of hadrosauroids. In relatively primitive ones, such asJinzhousaurusWang & Xu, 2001,EquijubusYou et al., 2003,XuwulongYou et al., 2011,ProbactrosaurusRozhdestvensky, 1966,EolambiaKirkland, 1998,ProtohadrosHead, 1998, andLevnesoviaSues & Averianov, 2009, the dentary teeth are large and shield-shaped relative to the alveolar trough (Norman, 2015: character 65). In contrary, inDatonglong, as well as inBactrosaurus,Gilmoreosaurus,Nanningosaurus,Zhanghenglong,Tethyshadros,Telmatosaurus, and hadrosaurids, the dentary teeth are miniaturized, and the lateral alveolar walls of tooth grooves became narrow and parallel-sided, rather than shaped by successional dentary crowns (Norman, 2015: character 66).ProbactrosaurusandLevnesoviapossess both large and shield-shaped teeth and narrow and parallel-sided lateral alveolar walls.Bactrosaurusis unique in having a height/width ratio greater than 3.1 as in hadrosaurids, de fi ning the dentary tooth crown in the mid-tooth row as an elongate lanceolate rather than diamond-shaped contour (Gilmore, 1933; Sues and Averianov, 2009; Prieto-Marquez, 2011).

The developments of ridges on the enameled lingual surface of the dentary tooth crown show various complex patterns and are characteristic for different taxa among hadrosauroids. The pattern inDatonglongshows a unique combination of character states. Generally, the positon of the primary ridge shifted from distal to median on the crown, and the number of ridges reduced from fi ve or more to only one primary ridge. In all hadrosaurids, the primary ridge is median placed; inZhanghenglong, the primary ridge is either median or slightly distally offset; while inDatonglongand all other known hadrosauroids, the primary ridge is distally offset (Xing et al., 2014: character 7). Therefore,Datonglongis probably more basal thanZhanghenglongbased on the placement of the primary ridge.

One the other hand,Datonglongpossesses two ridges as inTelmatosaurus, while many others includingProtohadros,Levnesovia,Bactrosaurus, andTethyshadroshave three or more (Xing et al., 2014: character 6), andGilmoreosaurus,EolambiaandClaosaurushave only oneridge (the primary ridge). Although both possess secondary ridge, it is strongly developed inDatonglong, in contrary to the faint one inTelmatosaurus. Therefore,Datonglongis probably more basal thanGilmoreosaurusbut more advanced thanBactrosaurusbased on the number of ridges on the dentary tooth crown.

The inclination of the coronoid process shifted from caudally-, vertically-, to rostrally inclined along the course of hadrosauroid evolution. InShuangmiaosaurusit is caudally inclined, inDatonglongit is basically vertical, while inNanningosaurusit is rostrally inclined. Therefore,Datonglongis probably more advanced thanShuangmiaosaurus, but less so thanNanningosaurusbased on the nature of the coronoid process. It is possible that as a non-hadrosaurid hadrosauroid,Nanningosaurusconvergently achieved a rostrally inclined coronoid process as in hadrosaurids.

Datonglongis unique in possessing two functional teeth in each alveolus at least along the mid- and caudal portion and probably along the entire occlusal plane. In primitive hadrosauroids, such asJinzhousaurus, only one functional tooth exists in each alveolus. In more advanced non-hadrosaurid hadrosauroids, such asEquijubus,Xuwulong,Probactrosaurus,Eolambia,Bactrosaurus,Gilmoreosaurus, andTethyshadros, a second functional tooth may occur in some alveoli in the middle portion of the dentary tooth row, inZhanghenglongone mid-alveolus possesses a third functional tooth although its rostral and caudal portions still having one, while inTelmatosaurus(Norman, 2015: character 54) and hadrosaurids, a third functional tooth adds on at least in some alveoli (Xing et al., 2014: character 4). Therefore,Datonglongseems to represent an advanced non-hadrosaurid hadrosauroid based on the development and pattern of functional teeth.

The above discussion shows a complex pattern in the evolution of dentary, especially dentary dentition among advanced non-hadrosaurid hadrosauroids. Derived hadrosaurid features occurred earlier frequently in non-hadrosaurids, such as high height/width ratio of tooth crown inBactrosaurus, one primary and one faint ridges inGilmoreosaurus, median placed primary ridge inZhanghenglong, rostrally inclined coronoid process inNanningosaurus, and two functional teeth per alveolus along at least mid- and caudal portion of the tooth row inDatonglong. This implies incredible diversities and attempts close to the origin of Hadrosauridae and how dif fi cult to elucidate their phylogenetic relationships.

Acknowledgments We are grateful to Mr. Tang Zhi-Lu for arranging this study; to the crew of the Shanxi Museum of Geology for discovering, excavating, and preparing the specimen; to reviewers for reviewing the ms. Funding was provided by the Hundred Talents Project of the Chinese Academy of Sciences, the National Natural Science Foundation of China (41472020), and the Department of Land and Resources of Shanxi Province.

Barrett P M, You H L, Upchurch P et al., 1998. A new ankylosaurian dinosaur (Ornithischia: Ankylosauria) from the Upper Cretaceous of Shanxi Province, People’s Republic of China. J Vert Paleont, 18: 376-384

Brett-Surman M K, 1979. Phylogeny and palaeobiogeography of hadrosaurian dinosaurs. Nature, 277: 560-562

Carrano M T, Benson R B J, Sampson S D, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). J Syst Palaeont, 10: 211-300

Chen Y G, Feng Z C, Fan Z Z, 1983. The age of the Huiquanbao Formation in the area of Yangyuan and Yuxian, Hebei. Region Geol China, 7: 39-45

Dalla Vecchia F M, 2009.Tethyshadros insularis, a new hadrosauroid dinosaur (Ornithischia) from the upper Cretaceous of Italy. J Vert Paleont, 29: 1100-1116

D’Emic M D, Mannion P D, Upchurch P et al., 2013. Osteology ofHuabeisaurus allocotus(Sauropoda: Titanosauriformes) from the Upper Cretaceous of China. PLoS ONE, 8: e69375

Dollo L, 1888. Iguanodontidae et Camptonotidae. C R Acad Paris, 106: 775-777

Gilmore C W, 1933. On the dinosaurian fauna of the Iren Dabasu Formation. Bull Am Mus Nat Hist, 67: 23-78

Godefroit P, Dong Z M, Bultynck P et al., 1998. Sino-Belgian Cooperation Program: ‘Cretaceous dinosaurs and mammalsfrom Inner Mongolia’ 1. NewBactrosaurus(Dinosauria: Hadrosauroidea) material from Iren Dabasu (Inner Mongolia, P. R. China). Bull Inst R Sci Nat Belg, 68: 3-70

Head J J, 1998. A new species of basal hadrosaurid (Dinosauria, Ornithischia) from the Cenomanian of Texas. J Vert Paleont, 18: 718-734

Kirkland J I, 1998. A new hadrosaurid from the Upper Cedar Mountain Formation (Albian-Cenomanian: Cretaceous) of eastern Utah - the oldest known hadrosaurid (Lambeosaurine?). In: Lucas S G, Kirkland J I, Estep J W eds. Lower and Middle Cretaceous Terrestrial Ecosystems,Vol. 14. New Mexico Museum of Natural History and Science Bulletin. Albuquerque: New Mexico Museum of Natural History and Science. 283-295

Liu X Q, 1983. A new dinosaur locality in Tianzhen County, Shanxi Province. Vert PalAsiat, 21(3): 272

Marsh O C, 1881. Principal characters of American Jurassic dinosaurs. Part V. Am J Sci, Ser 3, 21: 417-423

Marsh O C, 1890. New Cretaceous dinosaurs. Am J Sci, 39: 422-425

Mo J Y, Zhao Z R, Wang W et al., 2007. The fi rst hadrosaurid dinosaur from southern China. Acta Geol Sin-Engl Ed, 81: 550-554

Norman D B, 2015. On the history, osteology, and systematic position of the Wealden (Hastings group) dinosaurHypselospinus fi ttoni(Iguanodontia: Styracosterna). Zool J Linn Soc, 173(1): 92-189

Nopcsa F B, 1903.Telmatosaurus, new name for the dinosaurLimnosaurus. Geol Mag, 410: 94-95

Owen R, 1842. Report on British fossil reptiles. Rep Br Assoc Adv Sci, 9: 60-204

Pang Q Q, Cheng Z W, 1998. A new ankylosaur of Late Cretaceous from Tianzhen, Shanxi. Prog Nat Sci, 8(3): 326-334

Pang Q Q, Cheng Z W, 2000. A new family of sauropod dinosaur from the Upper Cretaceous of Tianzhen, Shanxi Province, China. Acta Geol Sin-Engl Ed, 74: 117-125

Pang Q Q, Cheng Z W, 2001. The Late Cretaceous dinosaur fauna and strata from Tianzhen, Shanxi and Yangyuan, Hebei, China. In: Deng T, Wang Y Q eds. Proceedings of the Eighth Annual Meeting of the Chinese Society of Vertebrate Paleontology. Beijing: China Ocean Press. 75-82

Pang Q Q, Cheng Z W, Yang J P et al., 1996. The preliminary report on Late Cretaceous dinosaur fauna expeditions in Tianzhen, Shanxi. J Hebei Coll Geol, 19(3-4): 227-235

Prieto-Marquez A, 2011. Cranial and appendicular ontogeny ofBactrosaurus johnsoni, a hadrosauroid dinosaur from the Late Cretaceous of northern China. Palaeontology, 54: 773-792

Prieto-Marquez A, Norell M A, 2010. Anatomy and relationships ofGilmoreosaurus mongoliensis(Dinosauria: Hadrosauroidea) from the Late Cretaceous of Central Asia. Am Mus Novit, 3694: 1-49

Rozhdestvensky A K, 1966. New iguanodonts from Central Asia. Int Geol Rev, 9: 556-566

Seeley H G, 1887. On the classification of the fossil animals commonly named Dinosauria. Proc R Soc London, 43: 165-171

Sereno P C, 1986. Phylogeny of the bird-hipped dinosaurs (Order Ornithischia). Natl Geogr Res, 2: 234-256

Sereno P C, 1997. The origin and evolution of dinosaurs. Annu Rev Earth Planet Sci, 25: 435-489

Sereno P C, 1998. A rationale for phylogenetic de fi nitions, with application to the higher-level taxonomy of Dinosauria. Neues Jahrb Geol Pal?ont Abh, 210: 41-83

Sereno P C, 2005. Stem Archosauria Version 1.0. TaxonSearch. Available at http://wwwtaxonsearchorg/Archive/stemarchosauria-10php, Accessed January 15, 2015

Sues H D, Averianov A, 2009. A new basal hadrosauroid dinosaur from the Late Cretaceous of Uzbekistan and the early radiation of duck-billed dinosaurs. Proc R Soc B-Biol Sci, 276: 2549-2555

Sullivan R M, 1999.Nodocephalosaurus kirtlandensis, gen. et sp. nov., a new ankylosaurid dinosaur (Ornithischia: Ankylosauria) from the Upper Cretaceous Kirtland Formation (Upper Campanian), San Juan Basin, New Mexico. J Vert Paleont, 19: 126-139

Sullivan R M, 2000. The taxonomic status ofShanxiatianzhenensis(Ornithischia, Ankylosauridae): a response to Sullivan (1999) - Reply. J Vert Paleont, 20: 218-219

Thompson R S, Parish J C, Maidment S C R et al., 2012. Phylogeny of the ankylosaurian dinosaurs (Ornithischia: Thyreophora). J Syst Palaeont, 10: 301-312

Tsogtbaatar K, Weishampel D B, Evans D C et al., 2014. A new hadrosauroid (Plesiohadrosdjadokhtaensis) from the Late Cretaceous Djadokhtan Fauna of southern Mongolia. In: Eberth D A, Evans D C eds. Hadrosaurs. Bloomington & Indianapolis: Indiana University Press. 108-135

Upchurch P, Barrett P M, 2000. The taxonomic status ofShanxia tianzhenensis(Ornithischia, Ankylosauridae): a response to Sullivan (1999). J Vert Paleont, 20: 216-217

Wang R F, You H L, Xu S C et al., 2013. A new hadrosauroid dinosaur from the early Late Cretaceous of Shanxi Province, China. PLoS ONE, 8(10): e77058

Wang X L, Xu X, 2001. A new iguanodontid (Jinzhousaurus yangigen. et sp. nov.) from the Yixian Formation of western Liaoning, China. Chin Sci Bull, 46(19): 1669-1672

Wiman C, 1929. Die Kriede-dinosaurier aus Shantung. Palaeont Sin, Ser C, 6: 1-67

Xing H, Wang D Y, Han F L et al., 2014. A new basal hadrosauroid dinosaur (Dinosauria: Ornithopoda) with transitional features from the Late Cretaceous of Henan Province, China. PLoS ONE, 9(6): e98821

Xu X, Zhao X J, Lü J C et al., 2000. A new iguanodontian from Sangping Formation of Neixiang, Henan and its stratigraphical implication. Vert PalAsiat, 38: 185-191

You H L, Ji Q, Li J L et al., 2003a. A new hadrosauroid dinosaur from the mid-Cretaceous of Liaoning, China. Acta Geol Sin-Engl Ed, 77: 148-154

You H L, Luo Z X, Shubin N H et al., 2003b. The earliest-known duck-billed dinosaur from deposits of late Early Cretaceous age in northwest China and hadrosaur evolution. Cretaceous Res, 24(3): 347-355

You H L, Li D Q, Liu W C, 2011. A new hadrosauriform dinosaur from the Early Cretaceous of Gansu Province, China. Acta Geol Sin-Engl Ed, 85: 51-57

山西天鎮(zhèn)晚白堊世一新鴨嘴龍超科恐龍

續(xù)世朝1尤海魯2*王嘉瑋3王鎖柱1伊 劍1賈 磊1,2

(1 山西地質(zhì)博物館 太原 030024)
(2 中國科學院脊椎動物演化與人類起源重點實驗室，中國科學院古脊椎動物與古人類研究所 北京 100044 * 通訊作者)
(3 俄羅斯羅蒙諾索夫莫斯科國立大學地質(zhì)系 莫斯科 119991)

記述了山西天鎮(zhèn)晚白堊世灰泉堡組發(fā)現(xiàn)的一鴨嘴龍超科恐龍新屬種：天鎮(zhèn)大同龍Datonglong tianzhenensisgen. et sp. nov.。標本為一保存較完好的右側齒骨并帶牙齒。與其他鴨嘴龍超科恐龍不同，大同龍每個齒槽有兩個功能齒，而且舌面齒冠上嵴的發(fā)育也具有獨特的特征組合(主嵴向遠中方向偏移；次嵴發(fā)育；無其他附嵴；主嵴上半段略后彎)。對比研究發(fā)現(xiàn)，在較進步的非鴨嘴龍科鴨嘴龍超科類群中齒骨和牙齒的演化過程復雜，許多鴨嘴龍科的特征在這些類群中時而出現(xiàn)，如：巴克龍Bactrosaurus齒冠具有較大的高/寬比；吉爾摩龍Gilmoreosaurus齒冠上只有一個主嵴和纖弱的附嵴；張衡龍Zhanghenglong齒冠上主嵴中置；南寧龍Nanningosaurus下頜冠狀突前傾；大同龍Datonglong每個齒槽有兩個功能齒。這表明為了獲得更有效的牙齒咀嚼方式，在鴨嘴龍科起源前它的姐妹群們曾進行過多種嘗試；這也告誡我們，受大量趨同演化的影響，要厘清這些類群間的關系并非易事。

山西天鎮(zhèn)，晚白堊世，恐龍，鴨嘴龍超科

Q915.864

A

1000-3118(2016)01-0067-12

2015-04-13

Xu S C, You H L, Wang J W et al., 2016. A new hadrosauroid dinosaur from the Late Cretaceous of Tianzhen, Shanxi Province, China. Vertebrata PalAsiatica, 54(1): 67-78

中國科學院百人計劃、國家自然科學基金(批準號：41472020)和山西省國土資源廳資助。