杨氏马门溪龙（Mamenchisaurus youngi）的骨骼特征与马门溪龙类的系统关系分类
|
摘要
中国最著名的蜥脚类恐龙——马门溪龙,因长期缺乏可靠的头骨及全面的描述,致使分类位置众说纷纭,影响了对整个蜥脚类系统发育历史的讨论。本文选择已知保存最好的马门溪龙标本——杨氏马门溪龙为研究对象,对其较完整的骨架,尤其是完好的头骨进行了详细描述。对比分析查明:马门溪龙具有圆顶龙类型的头骨,头后骨骼具有既不同于圆顶龙—腕龙类,也不同于梁龙—叉龙类的特性,故建议将马门溪龙及其相似的属种——峨眉龙、盘足龙等,作为蜥脚类一个单独的科级分类阶元对待,称马门溪龙科。该科以头骨轻巧、头骨孔发达、鼻孔侧位、牙齿勺状、下颌瘦长为特征;头后骨骼则以颈部特长,颈椎数目多,颈肋细长为主要特征;前部尾椎在早期属种中为平凹形,在后期属种中均为前凹形。
本文对较为罕见的蜥脚类恐龙的皮肤化石和巩膜环做了详细描述和报道,同时对杨氏马门溪龙的脑腔、齿腔还运用计算机辅助断层扫描技术(CT)进行了观察,发现其脑腔甚小,但脑腔中已出现较进步的结构分区。蜥脚类的牙齿替换具有连续性,新牙的生长与老牙的齿根吸收是同时进行的;齿根吸收越多的牙齿,其齿冠的磨蚀迹也越显著。杨氏马门溪龙牙齿的磨蚀迹还揭示,这种植食性恐龙的食料可能较为粗糙。马门溪龙类细长的颈肋使颈部强度增强,可是也限制了颈椎间的相对活动以及整个颈部的柔韧性,使颈部变得僵硬,故其长颈不大可能像长颈鹿一样伸向高处,其觅食的适宜高度估计不会超过肩部高度的2米。
本文提出了新的蜥脚类恐龙系统分类方案,建议以马门溪龙科与火山齿龙科、蜀龙科、圆顶龙科、腕龙科、叉龙科、巨龙科构成蜥脚类分类的基本框架。蜥脚类由进步的原蜥脚类通过早、中侏罗世的原始蜥脚类演化而来,其中马门溪龙类主要分布于东亚地区,在中侏罗世就已从原始蜥脚类中分化出来,并在侏罗纪晚期达到繁盛和演化的高峰,进入白垩纪后则走向衰亡。在众多蜥脚类中,马门溪龙与峨眉龙是最容易混淆的两个属,它们的最大差别在于前部尾椎形态。生活于中侏罗世的峨眉龙的前部尾椎为微弱双凹型或平凹型,生活于晚侏罗世的马门溪龙的前部尾椎为前凹型。按此标准,“长寿峨眉龙”、“釜溪峨眉龙”、“毛氏峨眉龙”均该归入马门溪龙。目前,除“井研马门溪龙”因材料混杂而命名可疑外,马门溪龙属的有效种共5个,即建设马门溪龙、杨氏马门溪龙、合川马门溪龙、安岳马门溪龙和中加马门溪龙。
Previous opinions on the phylogenetic position of Mamenchisaurus, the most famous sauropod from China, varied because of the lack of the reliable skull and its complete description. This disadvantage has seriously affected and postponed scientific understanding of the phylogenetic history of the entire Sauropoda. Fortunately, the specimen best preserved known up to date of mamenchisaurs, Mamenchisaurus youngi, turned to be available to this study, so that a detailed description of its complete skeleton framework, specially of its skull, has become feasible. The results indicate that Mamenchisaurus has a camarasaurid-type skull and characteristic postcranials, which are different not only from those of camarasaurids-branchiosaurids, but also from those of diplodocuds-dicraeosaurids. Thus, it is suggested that Mamenchisaurus and some similar long-necked genera, such as Omeisaurus and Euhelopus, should be recognized as a distinct sauropoda branch, Mamenchisauridae. The skull of this family is characterized by its lighter structure with large opening, spatulate crown and thin mandibular. The characteristics of its postcranial skeleton include a very long neck, counting 17-19 cervicls associated with long and striaght cervical ribs. The tail is shorter than the neck and the chevrons in the middle caudals are forked. The posterior cervical and anterior dorsal spines are not bifurcated and the anterior caudals are slightly amphicoelous or platycoelous in primitive forms, while in the advanced forms they have the bifurcated spines both on their posterior cervicals and anterior dorsals, and the precoelous centrum on their proximal caudals.Visual descriptions for the sclerotic ring and the fossilized skin of sauropods, which are rarely discovered, are presented here in this thesis. The computerized tomography (CT) scanning technology was used to investigate the cavity structures of the brain and tooth of the Mamenchisaurus youngi. The result displays that this dinosaur had a relatively small brain cavity (less than 80 ml), but with relatively advanced structural divisions, and that its teeth replacement was continuous in life. The absorption of an old tooth root was accompanied with the growth of a new tooth. The more the absorption amounts of the roots, the more prominent the worn facets on the crown. The developed worn facets of teeth also show that the food of
本文对较为罕见的蜥脚类恐龙的皮肤化石和巩膜环做了详细描述和报道,同时对杨氏马门溪龙的脑腔、齿腔还运用计算机辅助断层扫描技术(CT)进行了观察,发现其脑腔甚小,但脑腔中已出现较进步的结构分区。蜥脚类的牙齿替换具有连续性,新牙的生长与老牙的齿根吸收是同时进行的;齿根吸收越多的牙齿,其齿冠的磨蚀迹也越显著。杨氏马门溪龙牙齿的磨蚀迹还揭示,这种植食性恐龙的食料可能较为粗糙。马门溪龙类细长的颈肋使颈部强度增强,可是也限制了颈椎间的相对活动以及整个颈部的柔韧性,使颈部变得僵硬,故其长颈不大可能像长颈鹿一样伸向高处,其觅食的适宜高度估计不会超过肩部高度的2米。
本文提出了新的蜥脚类恐龙系统分类方案,建议以马门溪龙科与火山齿龙科、蜀龙科、圆顶龙科、腕龙科、叉龙科、巨龙科构成蜥脚类分类的基本框架。蜥脚类由进步的原蜥脚类通过早、中侏罗世的原始蜥脚类演化而来,其中马门溪龙类主要分布于东亚地区,在中侏罗世就已从原始蜥脚类中分化出来,并在侏罗纪晚期达到繁盛和演化的高峰,进入白垩纪后则走向衰亡。在众多蜥脚类中,马门溪龙与峨眉龙是最容易混淆的两个属,它们的最大差别在于前部尾椎形态。生活于中侏罗世的峨眉龙的前部尾椎为微弱双凹型或平凹型,生活于晚侏罗世的马门溪龙的前部尾椎为前凹型。按此标准,“长寿峨眉龙”、“釜溪峨眉龙”、“毛氏峨眉龙”均该归入马门溪龙。目前,除“井研马门溪龙”因材料混杂而命名可疑外,马门溪龙属的有效种共5个,即建设马门溪龙、杨氏马门溪龙、合川马门溪龙、安岳马门溪龙和中加马门溪龙。
Previous opinions on the phylogenetic position of Mamenchisaurus, the most famous sauropod from China, varied because of the lack of the reliable skull and its complete description. This disadvantage has seriously affected and postponed scientific understanding of the phylogenetic history of the entire Sauropoda. Fortunately, the specimen best preserved known up to date of mamenchisaurs, Mamenchisaurus youngi, turned to be available to this study, so that a detailed description of its complete skeleton framework, specially of its skull, has become feasible. The results indicate that Mamenchisaurus has a camarasaurid-type skull and characteristic postcranials, which are different not only from those of camarasaurids-branchiosaurids, but also from those of diplodocuds-dicraeosaurids. Thus, it is suggested that Mamenchisaurus and some similar long-necked genera, such as Omeisaurus and Euhelopus, should be recognized as a distinct sauropoda branch, Mamenchisauridae. The skull of this family is characterized by its lighter structure with large opening, spatulate crown and thin mandibular. The characteristics of its postcranial skeleton include a very long neck, counting 17-19 cervicls associated with long and striaght cervical ribs. The tail is shorter than the neck and the chevrons in the middle caudals are forked. The posterior cervical and anterior dorsal spines are not bifurcated and the anterior caudals are slightly amphicoelous or platycoelous in primitive forms, while in the advanced forms they have the bifurcated spines both on their posterior cervicals and anterior dorsals, and the precoelous centrum on their proximal caudals.Visual descriptions for the sclerotic ring and the fossilized skin of sauropods, which are rarely discovered, are presented here in this thesis. The computerized tomography (CT) scanning technology was used to investigate the cavity structures of the brain and tooth of the Mamenchisaurus youngi. The result displays that this dinosaur had a relatively small brain cavity (less than 80 ml), but with relatively advanced structural divisions, and that its teeth replacement was continuous in life. The absorption of an old tooth root was accompanied with the growth of a new tooth. The more the absorption amounts of the roots, the more prominent the worn facets on the crown. The developed worn facets of teeth also show that the food of
引文
马其鸿.四川盆地侏罗系—下白垩统瓣鳃类化石.见:四川盆地陆相中生代地层古生物编写组编著,四川盆地陆相中生代地层古生物,下篇,古生物论文集.成都:四川人民出版社,1982,582-622
王鸿祯等.中国古地理图集.北京:地图出版社,1-281
皮孝忠,欧阳辉,叶勇.四川自贡蜥脚类一新种.见:地质科学论文集.北京:中国经济出版社,1996,87-91
叶勇,欧阳辉,付乾明.四川自贡发现的合川马门溪龙新材料.古脊椎动物学报,2001,39(4):266-271
吕君昌,杜湘珂,朱奇志等.CT观察魏氏准葛尔翼龙(Dsungaripterus weii)头骨的脑腔形态.科学通报,1997,42(5):519-523
李奎.中国的蜥脚类恐龙化石及其层位.成都理工学院学报,1998,25(1):53-60
李奎,蔡开基.马门溪龙属(Mamenchisaurus)的分类位置及演化,成都理工学院学报,1997,24(2):102-107
李星学.中国地质时期的植物群.广州:广东科技出版社,1995,1-542
刘协章.四川盆地侏罗纪、白垩纪非海相瓣鳃类化石.见:四川盆地陆相中生代地层古生物编写组编著,四川盆地陆相中生代地层古生物,下篇,古生物论文集.成都:四川人民出版社,1982,529-581
沈炎彬,陈丕基.四川盆地侏罗、白垩纪叶肢介化石.见:四川盆地陆相中生代地层古生物编写组编著,四川盆地陆相中生代地层古生物,下篇,古生物论文集.成都:四川人民出版社,1982,392-415
宋之琛,尚玉珂.中国孢粉化石,第2卷,中生代孢粉.北京:科学出版社,2000,1-710
杨钟健.四川宜宾的一种新蜥脚类.古生物学报,1954,2(2):355-369
杨钟健,赵喜进.合川马门溪龙.见:中国科学院古脊椎动物与古人类研究所甲种专刊第8号.北京:科学出版社,1972,1-30
何信禄,李奎,蔡开基等.自贡大山铺峨眉龙—新种.成都地质学院学报,1984,增刊2:13-32
何信禄,李奎,蔡开基.蜥脚类(二)天府峨眉龙.见:四川自贡大山铺中侏罗世恐龙动物群(第四集)成都:四川科学技术出版社,1988,1-143
何信禄,杨绥华,蔡开基等.马门溪龙(蜥脚类恐龙)的新发现.见:地质科学论文集.北京:中国经济出版社,1996,83-86
张奕宏,杨代环,彭光照.四川自贡大山铺中侏罗世蜀龙新材料.成都地质学院学报,1984,增刊2:1-12
张奕宏.蜥脚类(一)李氏蜀龙.见:四川自贡大山铺中侏罗世恐龙动物群(第三集).成都:四川科学技术出版社,1988,1-67
张奕宏,李奎,曾清华.四川盆地晚侏罗世蜥脚类一新种.成都理工学院学报,1998,25(1):61-68
张玉光,李建军.井研马门溪龙(Mamenchisaurus jingyanensis)新材料的补充研究.见:第八届中国古脊椎动物学学术年会论文集.北京:海洋出版社,2001,35-39
张法奎,杜湘珂,罗德馨等.CT观察杨氏中国尖齿兽(Sinoconodon youngi)头骨化石标本的鼻部.古脊椎动物学报,1994,32(3):195-199
郑钟.蜀龙脑颅解剖.古脊椎动物学报,1991,29(4):108-118
欧阳辉.四川自贡大山铺蜥脚类—新属.自贡恐龙博物馆通讯,1989,第2期:10-14
欧阳辉,皮孝忠,叶勇.我国首例蜥脚类恐龙皮肤化石的发现.矿物岩石,1998增刊:146-149
唐烽,金幸生等.四川井研—完整的蜥脚类恐龙——毛氏峨眉龙.北京:海洋出版社,2001,1-128
候连海,周世武,曹幼枢.四川蜥脚类的新发现.古脊椎动物与古人类,1976,14(3):160-165
夏文杰等.自贡大山铺中侏罗世下沙溪庙组恐龙动物群的埋藏环境.成都地质学院学报,1984,增刊2:46-59
夏文杰等.恐龙埋藏环境与岩相古地理特征.成都:四川科学技术出版社,1988,1-88
夏宗实等.侏罗系.见:四川盆地陆相中生代地层古生物编写组编著,四川盆地陆相中生代地层古生物,上篇,地层.成都:四川人民出版社,1982,57-150
董枝明,唐治路.蜀龙动物群简报3蜥脚类.古脊椎动物与古人类,1984,22:69-75
董枝明,周世武,张奕宏.四川盆地侏罗纪恐龙化石.中国古生物志(新丙种),1983,第23号,1-145
董枝明,彭光照,黄大喜.蜥脚类骨质尾锤之发现.古脊椎动物学报,1989,27(3):219-224
魏明瑞,胡书生,张雁.云南禄丰原蜥脚类和蜥脚类恐龙食性研究.见:第八届中国古脊椎动物学学术年会论文集.北京:海洋出版社,2001,21-27
Barrett P M, Upchurch P, Sauropod feeding mechanisms: their bearing on palaeoecology. In: Ailing Sun and Yuangqing Wang (eds).Sixth Symposium on Mesozoic Terrestrial Ecosystems and Biota.China Ocean Press, Beijing, 1995, 107-110
Berman D S, Mcintosh J S, Skull and relationships of the Upper Jurassic sauropod Apatosaurus (Reptilia, Saurischia). Bulletin of the Carnegie Museum of Natural History. 1978, 8: 1-35
Bonaparte J F, Dinosaur: a Jurassic assemblage from Patagonia. Science, 1979, 205:1377-1379
Bonaparte J F, The early radiation and Phylogenetic relationships of Jurassic sauropod dinosaur, Based on vertebral anatomy. In: Padian K, ed. The Beginning of the Age of Dinosaurs. Cambridge: Cambridge University Press, 1986, 247-258
Borsuk-Bialynicka M, A new comarasaurid Opisthocoelicaudia gen. n. sp. n. from the Upper Cretaceous of Mongolia. Palaeontologica Polonica, 1977, 37: 5-64
Carroll R L, Vertebrate Paleontology and Evolution. W H Freeman and Company, New York, 1988, 698pp
Chapman R E, Shape analysis in the study of dinosaur morphology. In: Carpenter K, Currie P J , eds Dinosaur Systematics. Approaches and Perspectives. Cambridge: Cambridge University Press, 1990, 21-42
Conroy G C, Vannier M W, Noninvasive three-dimensional computer imaging of matrix-filled fossil skulls by high-resolution computed tomography. Science, 1984, 226: 456-458
Conroy G C, Dental development of the Taung skull from computerized tomography. Nature, 1987, 329: 625-627
Cooper M R, A reassessment of Vulcanodon karibaensis Raath (Dinosauria: Saurischia) and the origin of the sauropoda. Palaeontologica Africana 1984, 25:203-231
Currie P J, Zhao X J, A new troodontid (Dinosauria, Theropoda ) braincase from the Dinosaur Park Formation (Campanian ) of Alberta. Can. J. Earth Sci., 1993, 30: 2231-2247
Dong Z, Dinosaurian Faunas of China. Berlin: Springer-Verlag. 1992, 188pp
Dong Z, The dinosaur complexes of China and their biochronology. In: Ailing Sun and Yuangqing Wang (eds).Sixth Symposium on Mesozoic Terrestrial Ecosystems and Biota.China Ocean Press, Beijing, 1995,91-96
Forster C A, New information on the skull of Triceratops . Journal of Vertebrate Paleontology, 1996,16(2): 246-258
Galton P M, Basal Sauropodomorpha-Prosauropods. In: D B, Weishampel, P, Dodson and H, Osmolska (eds), The Dinosanria. University of California Press, Butkeley, 1990, 320-344
Gillette D D, Seismosaurus halli, gen. et sp. nov., a new sauropod dinosaur from the Morrison Formation ( Upper Jurassic/Lower Cretaceous ) of New Mexico, U.S.A.. Journal of Vertebrate Paleontology,
1991, 11(4): 417-433
Gilmore C W, A nearly complete articulated skeleton of Camarasaurus, a saurischian dinosaur from the Dinosaur National Monument, Utah. Memoris of the Carnegie Museum of Natural history 1925, 10: 347-384
Gilmore C W, Osteology of Apatosaurus. Memoris of the Carnegie Museum of Natural history, 1936, 11: 177-269
Gilmore C W, On a newly mounted skeletone of Diplodocus in the United States National Museum. Proceedings of the U.S. Natural Museum 1932, 81:1-21
Haubits B, Prokop M, Dohring W, et al., Computed tomography of Archaeopteryx. Paleobiology,1988, 14(2): 206-213
Hatcher J B, Osteology of Haplocanthosaurus with description of a new species, and remarks on the probable habits of the Sauropoda and the age and origin of the Atlantosaurus beds. Memoris of the Carnegie Museum of Natural history, 1903, 2: 1-72
Hollan W J, The ostelogy of Diplodocus Marsh. Memoris of the Carnegie Museum, 1906, 2: 225-278
Honsfield G N, Computerized transverse axial scanning tomography. British Journal of Radiology, 1973, 46:10-16
Hopson J A, Relative brain size in dinosaurs. In:R D K,Thomas and E C Olson (eds) ,A Cold Look at the Warm Blooded Dinosaurs. American Association for the Advancement of Science Symposium 28, 1977,287-310
Jerison H J Brain evolution and dinosaur brains. American Naturalist, 1968, 103: 575-588
JanenschW, Die Wirebelsaule der Gattung Dicraeosaurus. Palaeontographica, 1929 1(2): 37-133
Janensch W, Die Schadel der Sauropoden Branchisaurus, Barosaurus und Dicraesaurus, aus den Tendaguru-Schichten Deutsch-Ostafrikas. Palaeontographica, 1935-36, 2: 147-298
Janensch W, Die Wirebelsaule von Branchiosaurus brancai. Palaeontographica, 1950, 3: 27-93
Ledley R S, DiChiro G, Luessenhop J J, et al., Compuuterized transaxial X-ray tomography of the human body. Science, 1974, 186: 207-212
Madsen J H, Mclntosh J S, Berman D C, Skull and atlas-axis complex of the Upper Jurassic Sauropod Camarasaurus Cope (Reptilia: Saurischia). Bulletin of Carnegie Museum of Natural history, 1995, No.31:115pp
Mateer N J, Mclntosh J S, A new reconstruction of the skull of Euhelopus zdanskyi (Saurischia: Sauropoda). Bull. Geol. Inst. Univ. Uppsala, 1985,11: 125-132
McGowan C, Computer tomography reveals further details of Excalibosaurus, A putative ancestor for the swordfish-like ichthyosaur Eurhinosaurus. Journal of Vertebrate Paleontology, 1989,9: 269-281
McGowan C, Computed tomography confirms that Eurhinosaurus (Reptilia: Ichthyosauria) does have a tailbend. Can. J. Earth Sci.,1990,27:1541-1545
Mclntosh J S, Berman D S, Description of the palate and lower jaw of the sauropod dinosaur Dipolodocus (Reptilia: Saurischia) with remarks on the nature of the skull of Apatosaurus. Journal of paleontology 1975,49:187-199
Mclntosh J S, Sauropoda. In: D B, Weishampel, P, Dodson and H, Osmolska (eds). The Dinosanria. University of California Press, Butkeley, 1990, 345-401
Mclntosh J S, Specices determination in sauropod dinosaurs with tentative suggestions for their classification.In: Carpenter K, Currie P J , eds Dinosaur Systematics. Approaches and Perspectives. Cambridge: CambridgeUniversity Press, 1990, 53-70
Mclntosh J S, Miles C A, Cloward K C, Parker J R, A new nearly complete skeleton of Camarasaurus. Bulletin ofGunma Museum of Natural History, 1996, No.1: 87pp
Nowinski A, Nemegtosaurus mongoliensis n. gen., n. sp. (Sauropoda) from the uppermost Cretaceous of Mongolia.Palaeotologia Polonica, 1971, 25: 57-81
Owen R, A description of a portion of the skeleton of the Cetiosaurns, a gigantic extinct saurian repitile occurring in the oolitic formations of different portions of England. Proceedings of Geological Society of London. 1841,3: 457-462
Osborn H F, A skeleton of Dipolodocus. Memoirs of the American Museum of Natural History, 1899, 1: 191-214
Osborn H F, Mook C C, Camarasaurus, Amphicoelias and other saruopods of Cope. Memoirs of the American Museum of Natural History, 1921,3: 241-387
Osborn H F, Manus, sacrum, and caudals of Saurupoda. Bulletin of the American Museum of Natural History, 1904, 20; 181-190
Ostrom J H, Mclntosh J S, Marsh' s Dinosaur. Yale University Press, New Haven, 1966, 388pp
Raath M A, Fossile vertebrate studies in Rhodesia: A new dinosaur (Reptilia, Saurischia) from near the
Triassic-Jurassic boundary. Arnoldia, 1972, 5: 1-37
Romer A, S, Osteology of the Reptiles. University of Chicago Press, Chicago, 1956, 772pp.
Romer A, S, The Vertebrate Body (4th Edition). W B Saunders Company, Philadelphia, 1970, 601pp
Russell D A, The role of central Asia in dinosaurian biogeography. Can. J. Earth Sci., 1993, 30: 2002-2012
Russell S A, Zheng Z, A large mamenchisaurid from Junggar Basin, Xinjiang, People's Republic of china. Can. J. Earth Sci., 1993, 30: 2082-2095
Rothschild B M, Berman D S, Fusion of caudal vertebrae in late Jurassic sauropods. Journal of Vertebrate Paleontology, 1991,11(1):29-36
Salgado L, Calvo J O, Cranial osteology of Amargasaurus cazaui Salgado & Bonaparte (Sauropoda,Dicraeosauridae) from the Neocomian of Patagonia. Ameghiniana, 1992, 29(4): 337-346
Salgado L, Coria R A, Calvo J O, Evolution of titanosaurid sauropods. I: phylogenetic analysis based on the postcranial evidence. Ameghiniana, 34: 3-32
Sereno P C, The origin and evolution of dinosaurs. Annu. Rev. Earth Planet Sci. 1997,25: 435-489
Steel R, Saurischia. In O. Kuhn(eds): Encyclopeolia of paleoher petology. Stuttgart: Gustav Fischer Verlag, 1970, 87pp.
Sereno P C, Prosauropod monophyly and basal sauropodomorph phylogeny. Journal of Vertebrate Paleontology, 1990, 9(suppl.): 38A
Upchurch P, The phylogenetic relationships of sauropod dinosaurs.Zool J Linn Soc, 1998, 124: 43-103
White T E, The braincase of Camarasaurus lentus (Marsh). J. Pal., 1958, 32: 477-494
Wilson J A, Sereno P C, Higher-level phylogeny of sauropod dinosaurs. Jorunal of Vertebrate Paleontology, 1994, 14:42A
Wilson J A, Sereno P C, Early evolution and higher-level phyogeny of sauropod dinosaurs. Jorunal of Vertebrate Palaeotology, 1998, 18: 1-71
Wiman C, Die Kreide-Dinosaurier aus Shantung. Palaeontologia Sinica (c), 1929, 6: 1-67
Young C C, A new dinosaurian from Sinkiang. Palaeontologia Sinica, Series C, 1937, 2: 1-25
Young C C, On a new Sauropoda with notes on other Fragmentary Reptiles from Szechuan. Bull. Geol. Soc. China, 1939, Vol. XIX, No.3: 279-316
Young C C, New Sauropoda from China. Vertebrata PalAsiatica, 1958, Vol. II, No.l: 1-28
Yu C, Sauropod phylogeny: a preliminary cladistic analysis. Journal of Vertebrate paleontology.
王鸿祯等.中国古地理图集.北京:地图出版社,1-281
皮孝忠,欧阳辉,叶勇.四川自贡蜥脚类一新种.见:地质科学论文集.北京:中国经济出版社,1996,87-91
叶勇,欧阳辉,付乾明.四川自贡发现的合川马门溪龙新材料.古脊椎动物学报,2001,39(4):266-271
吕君昌,杜湘珂,朱奇志等.CT观察魏氏准葛尔翼龙(Dsungaripterus weii)头骨的脑腔形态.科学通报,1997,42(5):519-523
李奎.中国的蜥脚类恐龙化石及其层位.成都理工学院学报,1998,25(1):53-60
李奎,蔡开基.马门溪龙属(Mamenchisaurus)的分类位置及演化,成都理工学院学报,1997,24(2):102-107
李星学.中国地质时期的植物群.广州:广东科技出版社,1995,1-542
刘协章.四川盆地侏罗纪、白垩纪非海相瓣鳃类化石.见:四川盆地陆相中生代地层古生物编写组编著,四川盆地陆相中生代地层古生物,下篇,古生物论文集.成都:四川人民出版社,1982,529-581
沈炎彬,陈丕基.四川盆地侏罗、白垩纪叶肢介化石.见:四川盆地陆相中生代地层古生物编写组编著,四川盆地陆相中生代地层古生物,下篇,古生物论文集.成都:四川人民出版社,1982,392-415
宋之琛,尚玉珂.中国孢粉化石,第2卷,中生代孢粉.北京:科学出版社,2000,1-710
杨钟健.四川宜宾的一种新蜥脚类.古生物学报,1954,2(2):355-369
杨钟健,赵喜进.合川马门溪龙.见:中国科学院古脊椎动物与古人类研究所甲种专刊第8号.北京:科学出版社,1972,1-30
何信禄,李奎,蔡开基等.自贡大山铺峨眉龙—新种.成都地质学院学报,1984,增刊2:13-32
何信禄,李奎,蔡开基.蜥脚类(二)天府峨眉龙.见:四川自贡大山铺中侏罗世恐龙动物群(第四集)成都:四川科学技术出版社,1988,1-143
何信禄,杨绥华,蔡开基等.马门溪龙(蜥脚类恐龙)的新发现.见:地质科学论文集.北京:中国经济出版社,1996,83-86
张奕宏,杨代环,彭光照.四川自贡大山铺中侏罗世蜀龙新材料.成都地质学院学报,1984,增刊2:1-12
张奕宏.蜥脚类(一)李氏蜀龙.见:四川自贡大山铺中侏罗世恐龙动物群(第三集).成都:四川科学技术出版社,1988,1-67
张奕宏,李奎,曾清华.四川盆地晚侏罗世蜥脚类一新种.成都理工学院学报,1998,25(1):61-68
张玉光,李建军.井研马门溪龙(Mamenchisaurus jingyanensis)新材料的补充研究.见:第八届中国古脊椎动物学学术年会论文集.北京:海洋出版社,2001,35-39
张法奎,杜湘珂,罗德馨等.CT观察杨氏中国尖齿兽(Sinoconodon youngi)头骨化石标本的鼻部.古脊椎动物学报,1994,32(3):195-199
郑钟.蜀龙脑颅解剖.古脊椎动物学报,1991,29(4):108-118
欧阳辉.四川自贡大山铺蜥脚类—新属.自贡恐龙博物馆通讯,1989,第2期:10-14
欧阳辉,皮孝忠,叶勇.我国首例蜥脚类恐龙皮肤化石的发现.矿物岩石,1998增刊:146-149
唐烽,金幸生等.四川井研—完整的蜥脚类恐龙——毛氏峨眉龙.北京:海洋出版社,2001,1-128
候连海,周世武,曹幼枢.四川蜥脚类的新发现.古脊椎动物与古人类,1976,14(3):160-165
夏文杰等.自贡大山铺中侏罗世下沙溪庙组恐龙动物群的埋藏环境.成都地质学院学报,1984,增刊2:46-59
夏文杰等.恐龙埋藏环境与岩相古地理特征.成都:四川科学技术出版社,1988,1-88
夏宗实等.侏罗系.见:四川盆地陆相中生代地层古生物编写组编著,四川盆地陆相中生代地层古生物,上篇,地层.成都:四川人民出版社,1982,57-150
董枝明,唐治路.蜀龙动物群简报3蜥脚类.古脊椎动物与古人类,1984,22:69-75
董枝明,周世武,张奕宏.四川盆地侏罗纪恐龙化石.中国古生物志(新丙种),1983,第23号,1-145
董枝明,彭光照,黄大喜.蜥脚类骨质尾锤之发现.古脊椎动物学报,1989,27(3):219-224
魏明瑞,胡书生,张雁.云南禄丰原蜥脚类和蜥脚类恐龙食性研究.见:第八届中国古脊椎动物学学术年会论文集.北京:海洋出版社,2001,21-27
Barrett P M, Upchurch P, Sauropod feeding mechanisms: their bearing on palaeoecology. In: Ailing Sun and Yuangqing Wang (eds).Sixth Symposium on Mesozoic Terrestrial Ecosystems and Biota.China Ocean Press, Beijing, 1995, 107-110
Berman D S, Mcintosh J S, Skull and relationships of the Upper Jurassic sauropod Apatosaurus (Reptilia, Saurischia). Bulletin of the Carnegie Museum of Natural History. 1978, 8: 1-35
Bonaparte J F, Dinosaur: a Jurassic assemblage from Patagonia. Science, 1979, 205:1377-1379
Bonaparte J F, The early radiation and Phylogenetic relationships of Jurassic sauropod dinosaur, Based on vertebral anatomy. In: Padian K, ed. The Beginning of the Age of Dinosaurs. Cambridge: Cambridge University Press, 1986, 247-258
Borsuk-Bialynicka M, A new comarasaurid Opisthocoelicaudia gen. n. sp. n. from the Upper Cretaceous of Mongolia. Palaeontologica Polonica, 1977, 37: 5-64
Carroll R L, Vertebrate Paleontology and Evolution. W H Freeman and Company, New York, 1988, 698pp
Chapman R E, Shape analysis in the study of dinosaur morphology. In: Carpenter K, Currie P J , eds Dinosaur Systematics. Approaches and Perspectives. Cambridge: Cambridge University Press, 1990, 21-42
Conroy G C, Vannier M W, Noninvasive three-dimensional computer imaging of matrix-filled fossil skulls by high-resolution computed tomography. Science, 1984, 226: 456-458
Conroy G C, Dental development of the Taung skull from computerized tomography. Nature, 1987, 329: 625-627
Cooper M R, A reassessment of Vulcanodon karibaensis Raath (Dinosauria: Saurischia) and the origin of the sauropoda. Palaeontologica Africana 1984, 25:203-231
Currie P J, Zhao X J, A new troodontid (Dinosauria, Theropoda ) braincase from the Dinosaur Park Formation (Campanian ) of Alberta. Can. J. Earth Sci., 1993, 30: 2231-2247
Dong Z, Dinosaurian Faunas of China. Berlin: Springer-Verlag. 1992, 188pp
Dong Z, The dinosaur complexes of China and their biochronology. In: Ailing Sun and Yuangqing Wang (eds).Sixth Symposium on Mesozoic Terrestrial Ecosystems and Biota.China Ocean Press, Beijing, 1995,91-96
Forster C A, New information on the skull of Triceratops . Journal of Vertebrate Paleontology, 1996,16(2): 246-258
Galton P M, Basal Sauropodomorpha-Prosauropods. In: D B, Weishampel, P, Dodson and H, Osmolska (eds), The Dinosanria. University of California Press, Butkeley, 1990, 320-344
Gillette D D, Seismosaurus halli, gen. et sp. nov., a new sauropod dinosaur from the Morrison Formation ( Upper Jurassic/Lower Cretaceous ) of New Mexico, U.S.A.. Journal of Vertebrate Paleontology,
1991, 11(4): 417-433
Gilmore C W, A nearly complete articulated skeleton of Camarasaurus, a saurischian dinosaur from the Dinosaur National Monument, Utah. Memoris of the Carnegie Museum of Natural history 1925, 10: 347-384
Gilmore C W, Osteology of Apatosaurus. Memoris of the Carnegie Museum of Natural history, 1936, 11: 177-269
Gilmore C W, On a newly mounted skeletone of Diplodocus in the United States National Museum. Proceedings of the U.S. Natural Museum 1932, 81:1-21
Haubits B, Prokop M, Dohring W, et al., Computed tomography of Archaeopteryx. Paleobiology,1988, 14(2): 206-213
Hatcher J B, Osteology of Haplocanthosaurus with description of a new species, and remarks on the probable habits of the Sauropoda and the age and origin of the Atlantosaurus beds. Memoris of the Carnegie Museum of Natural history, 1903, 2: 1-72
Hollan W J, The ostelogy of Diplodocus Marsh. Memoris of the Carnegie Museum, 1906, 2: 225-278
Honsfield G N, Computerized transverse axial scanning tomography. British Journal of Radiology, 1973, 46:10-16
Hopson J A, Relative brain size in dinosaurs. In:R D K,Thomas and E C Olson (eds) ,A Cold Look at the Warm Blooded Dinosaurs. American Association for the Advancement of Science Symposium 28, 1977,287-310
Jerison H J Brain evolution and dinosaur brains. American Naturalist, 1968, 103: 575-588
JanenschW, Die Wirebelsaule der Gattung Dicraeosaurus. Palaeontographica, 1929 1(2): 37-133
Janensch W, Die Schadel der Sauropoden Branchisaurus, Barosaurus und Dicraesaurus, aus den Tendaguru-Schichten Deutsch-Ostafrikas. Palaeontographica, 1935-36, 2: 147-298
Janensch W, Die Wirebelsaule von Branchiosaurus brancai. Palaeontographica, 1950, 3: 27-93
Ledley R S, DiChiro G, Luessenhop J J, et al., Compuuterized transaxial X-ray tomography of the human body. Science, 1974, 186: 207-212
Madsen J H, Mclntosh J S, Berman D C, Skull and atlas-axis complex of the Upper Jurassic Sauropod Camarasaurus Cope (Reptilia: Saurischia). Bulletin of Carnegie Museum of Natural history, 1995, No.31:115pp
Mateer N J, Mclntosh J S, A new reconstruction of the skull of Euhelopus zdanskyi (Saurischia: Sauropoda). Bull. Geol. Inst. Univ. Uppsala, 1985,11: 125-132
McGowan C, Computer tomography reveals further details of Excalibosaurus, A putative ancestor for the swordfish-like ichthyosaur Eurhinosaurus. Journal of Vertebrate Paleontology, 1989,9: 269-281
McGowan C, Computed tomography confirms that Eurhinosaurus (Reptilia: Ichthyosauria) does have a tailbend. Can. J. Earth Sci.,1990,27:1541-1545
Mclntosh J S, Berman D S, Description of the palate and lower jaw of the sauropod dinosaur Dipolodocus (Reptilia: Saurischia) with remarks on the nature of the skull of Apatosaurus. Journal of paleontology 1975,49:187-199
Mclntosh J S, Sauropoda. In: D B, Weishampel, P, Dodson and H, Osmolska (eds). The Dinosanria. University of California Press, Butkeley, 1990, 345-401
Mclntosh J S, Specices determination in sauropod dinosaurs with tentative suggestions for their classification.In: Carpenter K, Currie P J , eds Dinosaur Systematics. Approaches and Perspectives. Cambridge: CambridgeUniversity Press, 1990, 53-70
Mclntosh J S, Miles C A, Cloward K C, Parker J R, A new nearly complete skeleton of Camarasaurus. Bulletin ofGunma Museum of Natural History, 1996, No.1: 87pp
Nowinski A, Nemegtosaurus mongoliensis n. gen., n. sp. (Sauropoda) from the uppermost Cretaceous of Mongolia.Palaeotologia Polonica, 1971, 25: 57-81
Owen R, A description of a portion of the skeleton of the Cetiosaurns, a gigantic extinct saurian repitile occurring in the oolitic formations of different portions of England. Proceedings of Geological Society of London. 1841,3: 457-462
Osborn H F, A skeleton of Dipolodocus. Memoirs of the American Museum of Natural History, 1899, 1: 191-214
Osborn H F, Mook C C, Camarasaurus, Amphicoelias and other saruopods of Cope. Memoirs of the American Museum of Natural History, 1921,3: 241-387
Osborn H F, Manus, sacrum, and caudals of Saurupoda. Bulletin of the American Museum of Natural History, 1904, 20; 181-190
Ostrom J H, Mclntosh J S, Marsh' s Dinosaur. Yale University Press, New Haven, 1966, 388pp
Raath M A, Fossile vertebrate studies in Rhodesia: A new dinosaur (Reptilia, Saurischia) from near the
Triassic-Jurassic boundary. Arnoldia, 1972, 5: 1-37
Romer A, S, Osteology of the Reptiles. University of Chicago Press, Chicago, 1956, 772pp.
Romer A, S, The Vertebrate Body (4th Edition). W B Saunders Company, Philadelphia, 1970, 601pp
Russell D A, The role of central Asia in dinosaurian biogeography. Can. J. Earth Sci., 1993, 30: 2002-2012
Russell S A, Zheng Z, A large mamenchisaurid from Junggar Basin, Xinjiang, People's Republic of china. Can. J. Earth Sci., 1993, 30: 2082-2095
Rothschild B M, Berman D S, Fusion of caudal vertebrae in late Jurassic sauropods. Journal of Vertebrate Paleontology, 1991,11(1):29-36
Salgado L, Calvo J O, Cranial osteology of Amargasaurus cazaui Salgado & Bonaparte (Sauropoda,Dicraeosauridae) from the Neocomian of Patagonia. Ameghiniana, 1992, 29(4): 337-346
Salgado L, Coria R A, Calvo J O, Evolution of titanosaurid sauropods. I: phylogenetic analysis based on the postcranial evidence. Ameghiniana, 34: 3-32
Sereno P C, The origin and evolution of dinosaurs. Annu. Rev. Earth Planet Sci. 1997,25: 435-489
Steel R, Saurischia. In O. Kuhn(eds): Encyclopeolia of paleoher petology. Stuttgart: Gustav Fischer Verlag, 1970, 87pp.
Sereno P C, Prosauropod monophyly and basal sauropodomorph phylogeny. Journal of Vertebrate Paleontology, 1990, 9(suppl.): 38A
Upchurch P, The phylogenetic relationships of sauropod dinosaurs.Zool J Linn Soc, 1998, 124: 43-103
White T E, The braincase of Camarasaurus lentus (Marsh). J. Pal., 1958, 32: 477-494
Wilson J A, Sereno P C, Higher-level phylogeny of sauropod dinosaurs. Jorunal of Vertebrate Paleontology, 1994, 14:42A
Wilson J A, Sereno P C, Early evolution and higher-level phyogeny of sauropod dinosaurs. Jorunal of Vertebrate Palaeotology, 1998, 18: 1-71
Wiman C, Die Kreide-Dinosaurier aus Shantung. Palaeontologia Sinica (c), 1929, 6: 1-67
Young C C, A new dinosaurian from Sinkiang. Palaeontologia Sinica, Series C, 1937, 2: 1-25
Young C C, On a new Sauropoda with notes on other Fragmentary Reptiles from Szechuan. Bull. Geol. Soc. China, 1939, Vol. XIX, No.3: 279-316
Young C C, New Sauropoda from China. Vertebrata PalAsiatica, 1958, Vol. II, No.l: 1-28
Yu C, Sauropod phylogeny: a preliminary cladistic analysis. Journal of Vertebrate paleontology.