葡萄園龍屬
葡萄园龙属 化石时期：白堊紀晚期
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| 葡萄園龍骨架模型 展示於法國埃斯佩拉扎恐龍博物館 | |
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： | 恐龍總目 Dinosauria |
| 目： | 蜥臀目 Saurischia |
| 亚目： | †蜥脚形亚目 Sauropodomorpha |
| 下目： | †蜥腳下目 Sauropoda |
| 演化支： | †泰坦巨龍類 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 |
| 模式種 | |
| 奧德河葡萄園龍 Ampelosaurus atacis Le Loeuff, 1995
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葡萄園龍（屬名：Ampelosaurus，發音：/ˌæmpɪloʊˈsɔːrəs/ AM-pi-loh-SOR-əs，意為「藤蔓蜥蜴」）是一屬泰坦巨龍類的蜥腳類恐龍，生存於白堊紀晚期的歐洲，化石發現於法國南部奥德河畔康帕涅附近的下紅泥灰層。模式種奧德河葡萄園龍（Ampelosaurus atacis）於1995年由讓·呂勒福（Jean Le Loeuff）命名、發表。另一個可能的未命名種，將葡萄園龍的生存年代延伸到白堊紀最末期的馬斯垂克階，約7000萬至6600萬年前。
如同多數蜥腳類，葡萄園龍有著典型的長頸子及長尾巴，背部長著由皮內成骨構成的鱗甲，身長約15公尺及體重約15公噸。有超過500塊骨頭被歸入葡萄園龍，其中包括腦殼，因此能有較深入的解剖學研究。牠是白堊紀晚期北半球少數幾種發現化石較多的泰坦巨龍類，且多虧近年來受到的關注，使葡萄園龍成為法國最著名的恐龍之一。
發現
葡萄園龍最初是在法國南部奧德省奧德河畔康帕涅市鎮附近發現的。出土於下紅泥灰層下部層位，地質年代屬於晚白堊世馬斯垂克階早期，約7000萬年前。這些沉積物顯示過去這裡曾是個佈滿河道的氾濫平原。[1]1995年，法國古生物學家讓·呂勒福（Jean Le Loeuff）首次敘述、命名了模式種奧德河葡萄園龍（Ampelosaurus atacis）。屬名源自古希臘語的αμπελος「藤蔓」，象徵著「葡萄園的蜥蜴」，取自發現地南邊附近、里莫氣泡酒（Blanquette de Limoux）產地的葡萄園。種名是奧德河的拉丁化atax，意為「來自奧德河的」。[1]
首批遺骸於1989年出土自一個屍骨層，產出包含大量肋骨、背椎、尾椎以及許多肢骨，但除了牙齒之外沒有發現其他頭骨材料，還發現四個不同尺寸和形狀的皮內成骨。這些化石分屬於許多隻不同個體。自1989年以來，在法國同一地區相繼發現了更多化石，包含相對完整的骨架與一些頭骨及下頜材料。[2]
後來在2001年，當初的奧德河畔康帕涅化石點由一匹新人所再次找到。他們經過長達13年的搜索，終於發現了一具幾乎完整的骨架，其中涵蓋更多原本正模標本所缺乏的骨骼部位。化石後來全部運往埃斯佩拉扎的恐龍博物館保存，前後耗時近10年來清修與裝設骨架。[3]
另一個未命名物種於2007年在西班牙馬德里－瓦倫西亞高速鐵路建設工程期間，在卡斯提爾-拉曼查村莊附近，屬於原本就盛產許多動物化石的維亞巴德拉席耶拉層，一個稱為羅韋科（Lo Hueco）地點發現。在幾個月有限的時間內，60多名古生物學家和100名工人合作展開了大規模緊急挖掘活動，救回數千個坎潘階晚期至馬斯垂克階早期的植物、無脊椎動物、脊椎動物化石標本，其中包含葡萄園龍未命名種。[4]
敘述
如同許多蜥腳類，葡萄園龍具有典型的長頸部和長尾部，背部還具有由25至28公分長的皮內成骨所構成的裝甲。發現了四個皮內成骨分別展現了三種不同的形態：板狀、球狀、棘刺狀。葡萄園龍估計身長約15至16公尺，體重約15000公斤。[1][5]對於奧德河葡萄園龍的了解來自大量零散、關連的骨骼及牙齒。牠是來自法國本土知名度最高的恐龍之一。[1]科里亞等人（2015）曾提出葡萄園龍可能是某種島嶼侏儒化的結果，因為體型明顯小於其他泰坦巨龍類祖先。[6]
骨骼
葡萄園龍是歐洲已知最完整的泰坦巨龍類蜥腳類之一。自1989年起，在法國南部貝里尤（Bellevue）地點發現超過500具骨骼，包含大量保存良好的材料，已被歸入奧德河葡萄園龍。在初始敘述論文中，只檢驗了一顆牙齒和一些脊椎。2005年呂勒福發表了一篇關於所有已知材料的完整敘述，使得葡萄園龍更深入的被了解。所有材料都被歸入模式種，就算不同的肢骨之間比例上有所差異，相信都只跟個體發育差異有關。[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]
鑑定特徵
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]
葡萄園龍未命名種
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]
神經解剖學
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]
分類
尾椎特徵和皮內成骨的存在證實葡萄園龍屬於岩盔龍類，這是一群先進的泰坦巨龍類，也包含阿拉莫龍、薩爾塔龍等知名物種。[9]許多其他研究人員曾將葡萄園龍列為泰坦巨龍類、[10]薩爾塔龍科[7]或泰坦巨龍科[1][6][11][12]也許目前尚未能確認葡萄園龍的確切分類位置，但牠具有許多典型特徵，足以判斷的確屬於泰坦巨龍類。[1] 最近迪耶茲迪亞茲（2018）的研究將葡萄園龍和其他坎潘階至馬斯垂克階的法國和西班牙泰坦巨龍類一同組成一個演化支，稱為細長龍亞科。其中葡萄園龍與吉普賽龍互成姊妹群；而整個演化支的位置則介於薩爾塔龍科、風神龍族、隆柯龍類之間。[13]
| 岩盔龍類 Lithostrotia |
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參考來源
- ^ 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.
