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Miodentosaurus Brevis Cheng Et Al., 2007 (Diapsida: Thalattosauria): Its Postcranial Skeleton and Phylogenetic Relationships

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Miodentosaurus Brevis Cheng Et Al., 2007 (Diapsida: Thalattosauria): Its Postcranial Skeleton and Phylogenetic Relationships 第 卷 第 期 古 脊 椎 动 物 学 报 47 摇 1 pp.1 - 20 年 月 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 2009 1 VERTEBRATA PALASIATICA 摇 figs.1 - 8 短吻贫齿龙( 双孔亚纲:海龙目) : 头后骨骼及系统关系研究1) , 吴肖春1 2 程延年2 佐藤环3 单希瑛2 加摇 拿大自然博物馆摇 渥太华 摇 (1 摇 ON K1P 6P4) 中国台湾自然科学博物馆 台中 (2 摇 40453) 日本东京学艺大学 东京 (3 摇 184- 8501) 摘要 短吻贫齿龙是根据一保存较完整的头骨和下颌标本建立的 标本采自贵州关岭地区三 : ; 叠系法郎组灰岩 对其头后骨骼进行了详细描述 短吻贫齿龙在肩带的乌喙骨和间锁骨及 。 。 腰带的肠骨显示了与众不同的特征 例如 紧靠肩臼后方乌喙骨有一明显的凹 间锁骨后半 , : , 部比前部要窄很多 及肠骨背突末端加宽等 头骨进一步细修表明 其中一些部位需要进行 , 。 , 重新记述 依据头后骨骼和头骨中的新信息 对短吻贫齿龙的特征作了修订 系统关系分析 。 , 。 表明 贫齿龙是个 其下颌宽大于长的反关节突和肱骨缩小的三角 胸大肌嵴表 , askeptosaurid, - 明它与产自瑞士和意大利的 Askeptosaurus 关系最近 。 关键词 贵州关岭 三叠纪 海龙目贫齿龙 头后骨骼 系统关系 : , , , , 中图法分类号 文献标识码 文章编号 :Q915. 864摇 :A摇 :1000- 3118(2009)01- 0001- 20 MIODENTOSAURUS BREVIS CHENG ET AL., 2007 (DIAPSIDA: THALATTOSAURIA): ITS POSTCRANIAL SKELETON AND PHYLOGENETIC RELATIONSHIPS 1,2* 2 3 2 CanadWianUMXuiseauom鄄Cohf uNnature 摇 CHENG Yen鄄Nien 摇 SATO Tamaki 摇 SHAN Hsi鄄Yin (1 Ottawa ON K1P 6P4, PO Box 3443 STN “D冶, Canada *Corresponding author: 摇 xcwu@ mus鄄nature. ca) Museum of Natural Science (2 Taichung 40453, 1, Kuan Chien RD, Taiwan, China) Tokyo Gakugei University (3 Tokyo 184- 8501, 4- 1- 1 Nukui鄄Kita鄄Machi, Koganei City, Japan) Abstract Miodentosaurus brevis 摇 was first established on the basis of the skull and mandible of a fairly preserved skeleton from the Triassic Falang Formation, Guanling area, Guizhou Province. The descrip鄄 M. brevis tion of the postcranial skeleton reveals that is also distinct in the morphology of its girdle ele鄄 ments, such as the coracoid with a small embayment just posterior to the glenoid, interclavicle becoming much narrower posteriorly than anteriorly, and the ilium having a dorsal blade with a expanded distal end. With a further preparation, some of the skull anatomy are redescribed. Based on new information from both skull and the postcranial skeleton, the diagnosis of the taxon is revised. A phylogenetic analy鄄 M. brevis Askeptosaurus sis suggests that is an askeptosauroid, closely related to from Switzerland and Italy on the basis of two unequivocal synapomorphies, a retroarticular process broader than long and a reduced deltopectoral crest in the humerus. Key words Miodentosaurus 摇 Guanling, Guizhou; Triassic; Thalattosauria, ; postcranial skeleton; phylogeny 中国科学院 国家外国专家局创新团队国际合作伙伴计划资助 1) 、 。 收稿日期 :2008- 07- 03 古 脊 椎 动 物 学 报 卷 摇 2 摇 摇 摇 摇 摇 摇 47 1摇 Introduction The holotype specimen (housed in MNaiotidoennatlosMaursuesumbreovfisNatural Science, with a catalogue number of NMNS 004727 / F003960) of Cheng et al., 2007 was collected from the Falang Formation (upper Middle to lower Upper Triassic) of Guanling area, Guizhou Province, China. Its skull and mandible was first described by Cheng et al. (2007) while its post鄄 cranial skeleton was still in the matrix. Now the latter is fully prepared and described here. Before the postcranial description, an anatomical revision of the skull and mandible is made on the basis of a further prepMa.rabtrieovnisusing Sand Bluster. With information drawn from the postcranial skeleton, the diagnosMis.obfrevis is revised and its phylogenetic relationships among thalattosaurians are ana鄄 lysed. is an askeptosauroid (see the text below) and the morphological comparison is madeNmeoasiinnlaysawuirtuhs thhoeantagxia of the Askeptosauroidea (sensu Liu and RieSpipneals,au2r0u0s5h)o.angi Zhou, 20W00ay(aZohsaouruins gYeieni & Zhou, 2000)( = Zhou, 2W0a0y0ao(sZahuorusinbeYlluins et al., 2000)), Zhou, 2000 (Zhou in Yin et al., 2000), and Zhou,2000 (Zhou in Yin et al., 2000) were originally described as pachypleuro鄄 sauroids on the basis of unprepared specimens. Our examination of the true specimens indicates that both genera are thalattosaurians. The original description of the two genera was not accurate or wrong in mNan.yhopalnacgeis; it is, for instance, obvious that both have only twMoiordatehnetorsathuarnusfibvreevsisacral vertebrae. (Gmr001, type specimen) appears to be similar to but clearly differs from the latter in that the neural spine of dorsal vertebrae is not constricted around their base andN.thheoahnugmi erus is nearly symmetrically incurved on both sides. The paratype skMul.l b(rGevmisr002) of bears a median ridge along the dorsal mid鄄line of the premaxillae as in but the neural sNp.inheoaonfgits dorsal vertebrae shows no constriction around the base. Any taxo鄄 nomic verification of has to wait for a full preparation of the specimens. WIt.isbepllousssible tWha.t gtheeeitype and paratype of the taxon could represent two different thalattosaurians. W. belalunsd are separately based on a specimen preserved in ventral view. The specimen of was not accessible but the relatively long symphysis of its maMnd.ibbrleevis(see fig. 2, 4 in Plate II in Zhou, 2000 (inW.Ygieneiet al., 2000)) is not comparabMle.tboretvhiast of . There is no doubt that the speci鄄 men of (Gmr004)Wi.sbneollturseferable to because its mandibular symphysis isWevaeynaorsealua鄄 rtiuvsely longer than that of . Again, the true taxonomic status of the two species of can be clarified until their specimens are fully prepared. 2摇 Description and comparison Revision of the skull and mandibular anatomy 摇 Fine preparation of the skull using Sand Bluster shows that some skull elements were not properly described or even inaccurately interpreted in Cheng et al. (2007), which are clarified here. 1) The maxillary鄄jugal relationship along the ven鄄 tral margin of the orbit was figured in the previous study as that the maxilla extends posteriorly me鄄 dial to the jugal (see fig. 2A in Cheng et al., 2007). This is not the case but the opposite is true (Fig. 1), as in other thalattosaurians. 2) The frontal resembles that of most thalattosaurians in pos鄄 sessing a posterolateral process, but it is very peculiar in that the bone forms the anterior margin of the pineal foramen (Fig. 1). This is unique among thalattosaurians. 3) The parietal is also very distinctive in having a slender anterolateral process that wraps the posterolateral process of the fron鄄 tal. Again, this is unique among thalattosaurians. 4) The ventral edge of the surangular of the man鄄 dible is not as straight as figured in the study of Cheng et al. (2007) but abruptly convex at the posterior third of the bone; in other words, the surangular suddenly broadens ventrally at the pMlaicoe鄄 dsoentthoasat uthrues sburreavnisgular / angular suture turns downward (Fig. 2). This appeTahrsaltaottboesaunruiqsuaelefxoarndrae . The surangular / angular suture also turns downward in (see fig. 2 in Nicholls,1999) but it is much weaker and more anteriorly positioned. 期 吴肖春等 短吻贫齿龙 双孔亚纲 海龙目 头后骨骼及系统关系研究 1 : ( : ): 摇 3 FigM. 1io摇dePnhtoostaou(ruAs)braenvdis outline of the photo (B) of the skull, mandible, and the first four cervical vertebrae of Cheng et al., 2007 in dorsal (the skull and right ramus of the mandible) and lateral (the left r关am节us窝of the mandible and the four cervica下ls)颌v收iew肌s窝 隅骨 Abbreviations关: 节af.骨articular fossa ; amf. 环ad椎duc神tor经m弓andibular fossa ; a环n.椎a间ngu椎la体r ; ar. a枢rti椎cular ; atar. 颈at肋lantal neural arch 颈椎 ; atic. atla齿nta骨l intercentrum 外鼻孔; ax. axis 额骨; cr. cervical rib孔 ; cv. cer舌vic骨al vertebra 轭骨; d. dentary ; en. external naris上隅骨外;侧f. 突frontal ; fo.上fo颌ra骨men ; hy. 鼻hy骨oid ; j. jugal ; lp. la环ter椎al齿pr突ocess of surang顶ula骨r ;前m环. 椎maxilla ; n. nasal松果孔; od. odontoid proce前ss颌of骨axis ; p. parietal 前颌;骨pa嵴t. proat鄄 las 后; p额fo骨. pineal foramen ; pm. premaxilla ; pmc. premaxillary crest ; pof. postfro狭nta长l 切口样;的p后ps.额s骨lit鄄l顶ike骨p缝ostfro表nt明al /原pa来rie上tal颞su孔tur的e,位in置dicating the position前o额f t骨he supratemporal f翼e鄄 n骨estra / , 第五颈椎后关; 节prf突. prefrontal 方骨; pt. pterygoid ; pz5.方l骨eft髁prezygapophysis of cerv方ic骨al 杆vertebra 5 反; 关q.节q突uadrate ; qcd. 上qu隅ad骨rate condyle ;夹q板s. 骨quadrate shaft 鳞骨; rap. retroarticular pr上oc颞ess骨 ; sa. surangu鼓lar膜附嵴 ; sp. splenial ; sq. squamosal ; st. supratemporal ; tyc. tympanic crest 古 脊 椎 动 物 学 报 卷 摇 4 摇 摇 摇 摇 摇 摇 47 Miodento鄄 sFaiug.ru2s摇 brPehvoisto (A) and outline (B) of the skull, mandible, and the first four cervical vertebrae of Cheng et al., 2007 in ventral (the skull and right ramus of the mandible), medial (the left ramus of the mandible), a基nd枕lat骨eral (the four cervical基s)蝶vie骨ws 内鼻孔 Abbreviations冠as状in骨Fig. 1 plus bo. basioccipital 齿骨; 联bs合. b面asisphenoid ;齿ch骨. 齿choana ; cn. coronoid齿骨齿窝; dsy. dentary symphysial facet 第五齿骨; d齿t. 窝dentary tooth ;外d翼ts. 骨dentary tooth socket 上翼骨 ; dts5. socket for dentary内to颈oth动5脉孔 ; ec. ectopterygoid 上;颌e骨p. 沟epi状pte槽rygoid ; ica. 前int关ern节al骨carotid artery 腭骨 ; mgr. groove鄄like tr前ou颌gh骨in齿maxillary ; par.副p枕rea突rticular ; pl. palatine 方; 骨pm翼t. 骨pr支emaxillary tooth 翼; 骨pop间. 窝parocci鄄 pital process ; prq. 翼pte骨ry方goi骨d 支ramus of quadrate 眶;下p孔tv. pterygoid vacuity ; qrp. q夹ua板dra骨te联ra合mu关s o节f p面terygoid 镫骨 ; sof. suborbital fenestra ; sp上sy颞. 孔spl遗en迹ial symphysial f犁ac骨et ; sta. stapes ; stf. trace of supratemporal fenestra ; vo. vomer 期 吴肖春等 短吻贫齿龙 双孔亚纲 海龙目 头后骨骼及系统关系研究 1 : ( : ): 摇 5 Miodentosaurus brevis Fig. 3摇 Skeleton of Cheng et al., 2007 in ventral ( the skull and right ramus of the mandible), medial (the left ramus of the mandible), and lateral (the postcranial skeleton) views (circled areas are reconstructed) 古 脊 椎 动 物 学 报 卷 摇 6 摇 摇 摇 摇 摇 摇 47 Axial skeleton Miodentosaurus brevis 摇 With the skull and mandible, the postcranial skeleton of is articulated and well preserved although the zeugopodium, the manus of the left hind鄄 limb, and some caudal vertebrae (3- 5, 12, 13, about 20 of the posterior most section) are missing (Fig. 3). There are 87 vertebrae preserved, consisting of 38 presacrals (13
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