葡萄园龙属
葡萄园龙属 化石时期：白垩纪晚期
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| 葡萄园龙骨架模型 展示于法国埃斯佩拉扎恐龙博物馆 | |
科学分类 
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| 界： | 动物界 Animalia |
| 门： | 脊索动物门 Chordata |
| 纲： | 蜥形纲 Sauropsida |
| 总目： | 恐龙总目 Dinosauria |
| 目： | 蜥臀目 Saurischia |
| 亚目： | †蜥脚形亚目 Sauropodomorpha |
| 下目： | †蜥脚下目 Sauropoda |
| 演化支： | †泰坦巨龙类 Titanosauria |
| 演化支： | †岩盔龙类 Lithostrotia |
| 科： | †细长龙亚科 Lirainosaurinae |
| 属： | †葡萄园龙属 Ampelosaurus Le Loeuff, 1995 |
| 模式种 | |
| 奥德河葡萄园龙 Ampelosaurus atacis Le Loeuff, 1995
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葡萄园龙（属名：Ampelosaurus，发音：/ˌæmpɪloʊˈsɔːrəs/ AM-pi-loh-SOR-əs，意为“藤蔓蜥蜴”）是一属泰坦巨龙类的蜥脚类恐龙，生存于白垩纪晚期的欧洲，化石发现于法国南部奥德河畔康帕涅附近的下红泥灰层。模式种奥德河葡萄园龙（Ampelosaurus atacis）于1995年由让·吕勒福（Jean Le Loeuff）命名、发表。另一个可能的未命名种，将葡萄园龙的生存年代延伸到白垩纪最末期的马斯特里赫特阶，约7000万至6600万年前。
如同多数蜥脚类，葡萄园龙有着典型的长颈子及长尾巴，背部长著由皮内成骨构成的鳞甲，身长约15米及体重约15公吨。有超过500块骨头被归入葡萄园龙，其中包括脑壳，因此能有较深入的解剖学研究。牠是白垩纪晚期北半球少数几种发现化石较多的泰坦巨龙类，且多亏近年来受到的关注，使葡萄园龙成为法国最著名的恐龙之一。
发现
葡萄园龙最初是在法国南部奥德省奥德河畔康帕涅市镇附近发现的。出土于下红泥灰层下部层位，地质年代属于晚白垩世马斯特里赫特阶早期，约7000万年前。这些沉积物显示过去这里曾是个布满河道的泛滥平原。[1]1995年，法国古生物学家让·吕勒福（Jean Le Loeuff）首次叙述、命名了模式种奥德河葡萄园龙（Ampelosaurus atacis）。属名源自古希腊语的αμπελος“藤蔓”，象征着“葡萄园的蜥蜴”，取自发现地南边附近、里莫气泡酒（Blanquette de Limoux）产地的葡萄园。种名是奥德河的拉丁化atax，意为“来自奥德河的”。[1]
首批遗骸于1989年出土自一个尸骨层，产出包含大量肋骨、背椎、尾椎以及许多肢骨，但除了牙齿之外没有发现其他头骨材料，还发现四个不同尺寸和形状的皮内成骨。这些化石分属于许多只不同个体。自1989年以来，在法国同一地区相继发现了更多化石，包含相对完整的骨架与一些头骨及下颌材料。[2]
后来在2001年，当初的奥德河畔康帕涅化石点由一匹新人所再次找到。他们经过长达13年的搜索，终于发现了一具几乎完整的骨架，其中涵盖更多原本正模标本所缺乏的骨骼部位。化石后来全部运往埃斯佩拉扎的恐龙博物馆保存，前后耗时近10年来清修与装设骨架。[3]
另一个未命名物种于2007年在西班牙马德里－巴伦西亚高速铁路建设工程期间，在卡斯提尔-拉曼查村庄附近，属于原本就盛产许多动物化石的维亚巴德拉席耶拉层，一个称为罗韦科（Lo Hueco）地点发现。在几个月有限的时间内，60多名古生物学家和100名工人合作展开了大规模紧急挖掘活动，救回数千个坎潘阶晚期至马斯特里赫特阶早期的植物、无脊椎动物、脊椎动物化石标本，其中包含葡萄园龙未命名种。[4]
叙述
如同许多蜥脚类，葡萄园龙具有典型的长颈部和长尾部，背部还具有由25至28公分长的皮内成骨所构成的装甲。发现了四个皮内成骨分别展现了三种不同的形态：板状、球状、棘刺状。葡萄园龙估计身长约15至16米，体重约15000公斤。[1][5]对于奥德河葡萄园龙的了解来自大量零散、关连的骨骼及牙齿。牠是来自法国本土知名度最高的恐龙之一。[1]科里亚等人（2015）曾提出葡萄园龙可能是某种岛屿侏儒化的结果，因为体型明显小于其他泰坦巨龙类祖先。[6]
骨骼
葡萄园龙是欧洲已知最完整的泰坦巨龙类蜥脚类之一。自1989年起，在法国南部贝里尤（Bellevue）地点发现超过500具骨骼，包含大量保存良好的材料，已被归入奥德河葡萄园龙。在初始叙述论文中，只检验了一颗牙齿和一些脊椎。2005年吕勒福发表了一篇关于所有已知材料的完整叙述，使得葡萄园龙更深入的被了解。所有材料都被归入模式种，就算不同的肢骨之间比例上有所差异，相信都只跟个体发育差异有关。[2]
A. atacis is known from a few, well-preserved teeth and some cranial material, a tooth of which was described in its original description. The teeth differentiate Ampelosaurus from Magyarosaurus and Lirainosaurus, both of which are also from Europe. The teeth are different from the two later genera as Ampelosaurus has teeth that are roughly cylindrical in the top with thin expansions at the bottom. They are 21毫米（0.83英寸） high and about 6毫米（0.24英寸） wide. Those expansions give its teeth a slight constriction between the root and the crown. The teeth are also quite different from the peg-like teeth of titanosaurids.[2] The teeth, have a general morphology similar to titanosaurians Antarctosaurus and a braincase from Dongargaon. Another skull was described from southern France, but showed some differences, meaning there could have been at least two different titanosaurians in France during the Late Cretaceous.[2]
The scapula was found attached to a coracoid. In the longer direction, the bone is 72厘米（28英寸） long. The blade of the scapula, contrary to titanosaurs, is triangular, larger closer to the base. The blade narrows distally instead of showing an expansion. Differing from Magyarosaurus and Saltasaurus, the scapula does not have the dorsal crest at the base of the scapular blade. The shape of the coracoid is quadrangular,[1] and the coracoid has a thickened cranial margin.[2]
Ampelosaurus is also known from a pubis, about 75厘米（30英寸） long. It has a strong distal expansion, that is similar to a titanosaur from Brazil, and a large foramen. Contrary to the South American titanosaurs, the ilium does not have an expansion on the side. The ischium, known from a shaft, is unexpanded distally and very well developed. It is also very flat.[2] An incomplete radius is also known. It lacks the proximal and distal ends, and possesses a very prominent ridge along it.[2] The largest and best-preserved specimens are both ulnas. The right ulna has a total length of 395毫米（15.6英寸）. The right ulna is small and slender, is missing an olecranon, and has a well-marked ridge. The left ulna is larger, with a length of 725毫米（28.5英寸）.[2] The ulna has a deep radial fossa.[7]
About 27 femora are known from Bellevue, making them the most common bone, most of them more-or-less complete. They are very flat craniocaudally but otherwise, lack any unusual features.[2] In a study of its bone physiology, it was found that the maximum known femur length was 1,100毫米（43英寸）, which is much larger than previously known (802毫米（31.6英寸）). That means, that the ancestors Ampelosaurus were slightly smaller than itself.[8] 18 humeri are also known,[2] but none of them approach the size of the largest femur. The humeri do not have a free medullary cavity.[8] The humeri are robust, about 63厘米（25英寸） long, with very expanded distal and proximal ends.[2]
Three different types of osteoderms have been recovered at Bellevue. Two with large spines have been uncovered. At the base of each spine, which is about 12厘米（4.7英寸） tall, are two large foramina. The internal faces are all concavo-convex and have a ridge opposite the spine. In side view, they have two sections, a low half that progressively thickens, and a spine.[2]
The holotype specimen of A. atacis is a group of three connected dorsal vertebrae. Cervical vertebrae are uncommon from Bellevue and most of them are poorly preserved.[2] The vertebrae are opisthocoelous, with centra that lengthen toward the rear. Neural spines have been preserved on the vertebrae, and they face toward the rear. Two dorsal vertebrae were preserved with a complete, but very crushed neural arch, a neural spine with a characteristic shape: it is very widened distally and narrows downwards. The vertebrae also have an internal structure that is spongy, with very large cells.[1] The caudal vertebrae are all strongly procoelous and are compressed on the sides. The neural spines of the caudal vertebrae are very narrow and very tall and point toward the rear. The middle caudal vertebrae are more compressed on the sides that the others. Also, the centrum of middle caudal vertebrae is longer proportionally.[1] Two sacral vertebrae were known from Bellevue. They are poorly preserved, and are thought to belong to a juvenile. A complete sacrum was also found in 2002.[2]
鉴定特征
This set of characters was identified by Le Loeuff in his description of Ampelosaurus to distinguish it from all other genera: teeth that weakly spatulate; a laterally widening distal extremity of neural spines on the dorsal vertebrae; neural arch of the dorsal vertebrae inclining strongly towards the rear; the lack of a distal expansion on the scapular blade; the presence of a light, ventral crest on top of the scapula; the presence of plate, bulb, and spine shaped osteoderms;[1] and, in 2005, Le Loeuff added that the constriction of the neural spine on the dorsal and cervical vertebrae was also probably a characteristic of Ampelosaurus.[2]
葡萄园龙未命名种
A fossil braincase from Lo Hueco was tentatively assigned to an unnamed species of Ampelosaurus, A. sp., in a 2013 article in the journal PLoS ONE. The braincase was found to share many features with A. atacis, such as a back of the skull that is flat. The braincase, MCCM-HUE-8741, is small in size overall, with a front-to-back length of 100.8毫米（3.97英寸）, and the maximum width of the left half being 64.3毫米（2.53英寸）. Parts of the bottom half of the braincase are missing. Even though section are missing, the specimen does not appear to have been deformed much, as the left and right halves are not very different.[4]
Two frontals are preserved. They are each 57.3毫米（2.26英寸） long and 64.3毫米（2.53英寸） wide. The upper surface of each frontal is not smooth. One crest runs along the each frontal, and the together the two crests make up the orbital roof. Both parietals have also been found. The connection between them is marked by a ω-shaped crest. Viewed from the side, the parietal has two extensions. These extensions are not fully preserved, but they would have been on the border of the upper temporal fenestrae in their middle. Each parietal was preserved as 79.6毫米（3.13英寸） wide. The basioccipital of the specimen from Lo Hueco is unique as it has an occipital condyle that is much wider than tall. The occipital condyle has an irregular surface that was probably caused by the loss of the original cartilaginous covering. The complete braincase was especially low in the skull, and was oriented to the side. The occipital condyle is 28.6毫米（1.13英寸） wide and 15.8毫米（0.62英寸） tall.[4] The braincase floor is made by the parabasisphenoid. The prootic is a tall but not long bone. The basisphenoid is mostly on the side of it, along with the laterosphenoid, the parietal, and the otoccipital. The length of the prootic from the front to the back is around 10.6毫米（0.42英寸）.[4]
神经解剖学
Compared with Giraffititan, the inner ear of A. sp. shows a more basal morphology. That feature is possibly related to a restricted range of possible movements that involve head-turning.[4]
Like in Jainosaurus and most other non-avian archosaurs, the hindbrain and midbrain of A. sp. is relatively poorly preserved in the endocast. In contrast with TMM 40435 and a few other taxa such as cf. Cetiosaurus oxoniensis and Giraffatitan, no characteristic "nub" of the cerebellum can be seen. As in TMM 40435 and many other archosaurs, the back of the brain is especially narrow in A. sp.[4]
The cerebral region of the brain is separated from the rest of the brain by a distinct compression caused in the endocranial cavity. The rearmost part of the cerebral region of the braincase has a top with a small expansion. This is different from Jainosaurus. However, relatively much larger expansions are known in the diplodocoid sauropods Dicraeosaurus and Diplodocus. In MCCM-HUE-8741, the small opening in the skull roof middle is responsible for a swelling on the endocast that is suggestive of a pineal system. It is in the exact position where the pineal gland is expected to have been, between the forebrain and the midbrain.[4]
The semicircular canals are contracted, and they are highly curved. The semicircular system of MCCM-HUE-8741 shows also a basal morphology, because the semicircular canals do not attach to each other.[4]
分类
尾椎特征和皮内成骨的存在证实葡萄园龙属于岩盔龙类，这是一群先进的泰坦巨龙类，也包含阿拉莫龙、萨尔塔龙等知名物种。[9]许多其他研究人员曾将葡萄园龙列为泰坦巨龙类、[10]萨尔塔龙科[7]或泰坦巨龙科[1][6][11][12]也许目前尚未能确认葡萄园龙的确切分类位置，但牠具有许多典型特征，足以判断的确属于泰坦巨龙类。[1] 最近迪耶兹迪亚兹（2018）的研究将葡萄园龙和其他坎潘阶至马斯特里赫特阶的法国和西班牙泰坦巨龙类一同组成一个演化支，称为细长龙亚科。其中葡萄园龙与吉普赛龙互成姊妹群；而整个演化支的位置则介于萨尔塔龙科、风神龙族、隆柯龙类之间。[13]
| 岩盔龙类 Lithostrotia |
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参考来源
- ^ 1.00 1.01 1.02 1.03 1.04 1.05 1.06 1.07 1.08 1.09 Le Loeuff, J. Ampelosaurus atacis (nov. gen., nov. sp.), un nouveau Titanosauridae (Dinosauria, Sauropoda) du Crétacé supérieur de la Haute Vallée de l'Aude (France) (PDF). Comptes Rendus de l'Académie des Sciences. IIa. 1995, 321: 693–699.
- ^ 2.00 2.01 2.02 2.03 2.04 2.05 2.06 2.07 2.08 2.09 2.10 2.11 2.12 2.13 2.14 Le Loeuff, J. Osteology of Ampelosaurus atacis (Titanosauria) from Southern France. Tidwell, V.; Carpenter, K. (编). Thunder-Lizards: The Sauropodomorph Dinosaurs. Bloomington: Indiana University Press. 2005: 115–137. ISBN 978-0-253-34542-4.
- ^ Souillat, C. & Le Loeuff, J. Numéro Spécial 2002. La Lettre de Dinosauria. 2002: 1–23.
- ^ 4.0 4.1 4.2 4.3 4.4 4.5 4.6 4.7 Knoll, F.; Ridgely, R. C.; Ortega, F.; Sanz, J. L.; Witmer, L. M. Butler, Richard J , 编. Neurocranial Osteology and Neuroanatomy of a Late Cretaceous Titanosaurian Sauropod from Spain (Ampelosaurus sp.). PLOS ONE. 2013, 8 (1): e54991. Bibcode:2013PLoSO...854991K. PMC 3552955
. PMID 23355905. doi:10.1371/journal.pone.0054991 .
- ^ Paul, G.S., 2010 The Princeton Field Guide to Dinosaurs, Princeton University Press p. 207
- ^ 6.0 6.1 Salgado, L.; Coria, R.A. Sauropods of Patagonia: systematic update and notes on global sauropod evolution. Carpenter, K.; Tidwell, V. (编). Thunder-Lizards: The Sauropodomorph Dinosaurs. Bloomington: Indiana University Press. 2005: 430–453. ISBN 978-0-253-34542-4.
- ^ 7.0 7.1 Wilson, J.A. Sauropod dinosaur phylogeny: critique and cladistic analysis (PDF). Zoological Journal of the Linnean Society. 2002, 136 (2): 217–276. doi:10.1046/j.1096-3642.2002.00029.x .
- ^ 8.0 8.1 Klein, N.; Sander, P. M.; Stein, K.; Le Loeuff, J.; Carballido, J. L.; Buffetaut, E. Farke, Andrew A , 编. Modified Laminar Bone in Ampelosaurus atacis and Other Titanosaurs (Sauropoda): Implications for Life History and Physiology. PLOS ONE. 2012, 7 (5): e36907. Bibcode:2012PLoSO...736907K. PMC 3353997 . PMID 22615842. doi:10.1371/journal.pone.0036907 .
- ^ Weishampel, D.B.; Dodson, P.; Osmolska, H. The Dinosauria 2nd. Berkeley: University of California Press. 2004: 259–322, 588–593. ISBN 978-0-520-24209-8.
- ^ Wilson, J.A.; Sereno, P.C. Early evolution and higher-level phylogeny of sauropod dinosaurs. Society of Vertebrate Paleontology Memoir 5. Journal of Vertebrate Paleontology. 1998, 18: 1–68. doi:10.1080/02724634.1998.10011115.
- ^ Le Loeuff, J. Les dinosaures du Crétacé supérieur de l'Île Ibéro-Armoricaine [The dinosaurs of the Upper Cretaceous of the Ibero-Armorican Island]. de Carvalho, A.M.G.; Andrade, A.M.; dos Santos, V.F.; Cascalho, J.; Taborda, R. (编). I Encontro Internacional sobre Paleobiologia dos Dinossáurios. Lisboa: Museu Nacional de História Natural. 1998: 49–64.
- ^ Le Loeuff, J.; Buffetaut, E. Ampelosaurus atacis, the dinosaur of the wineyard. The Dinosaur Society UK Quarterly. 1996, 1 (4): 1–2.
- ^ Díez Díaz, V.; Garcia, G.; Pereda-Suberbiola, X.; Jentgen-Ceschino, B.; Stein, K.; Godefroit, P.; Valentin, X. The titanosaurian dinosaur Atsinganosaurus velauciensis (Sauropoda) from the Upper Cretaceous of southern France: New material, phylogenetic affinities, and palaeobiogeographical implications. Cretaceous Research. 2018, 91: 429–456. doi:10.1016/j.cretres.2018.06.015.
