# 原蜥腳下目[编辑]

| †原蜥脚下目 化石时期：三疊紀晚期到侏儸紀早期 |
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| 黃氏雲南龍
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| 保护状况
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| 无效状况 |
| 科学分类  |
| 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： | **†原蜥脚下目 Prosauropoda** von Huene, 1920 |
| 科
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| * 里奧哈龍科 Riojasauridae
* 板龍科 Plateosauridae
* 大椎龍科 Massospondylidae
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**原蜥腳下目**（Prosauropoda）或稱**原蜥腳類**，是群早期植食性恐龍，生存於三疊紀晚期到侏儸紀早期。牠們在三疊紀的時候，通常是所處環境的優勢植食性動物，並快速地演化為大體型，身長6到10公尺；但是到了侏羅紀被真蜥腳亞目所完全取代、並且滅亡。所有原蜥腳類恐龍都有小型頭部、長頸部、前肢較後肢短、以及非常大的拇指尖爪（從槽齒龍繼承而來）可用來防衛。大部分原蜥腳類恐龍是半二足動物，但至少有一大型物種（里奧哈龍）是完全四足動物。

## 不斷改變的定義

原蜥腳下目起初被定義為二足的三疊紀恐龍，是體型巨大的蜥腳下目恐龍的早期祖先。2003年的親緣分支分類法研究顯示原蜥腳下目是個獨立演化支，而非蜥腳下目的祖先。2004年對於大椎龍的研究顯示原蜥腳下目的確是單系群。這群恐龍是蜥腳下目的姐妹分類單元，而非直系祖先。但是近期研究表示原蜥腳下目是個并系群。

問題在於哪些屬被歸類於原蜥腳下目。在1997年，保羅·阿普徹奇（Paul Upchurch）提出一個以某節點為基礎的定義，包含：貝里肯龍科、槽齒龍科、近蜥龍科、板龍科、黑丘龍科、以及蜥腳形亞目中親緣關係接近原蜥腳類，而離蜥腳類較遠的所有物種。最近，亞當·耶茨（Adam Yates）以對於早期蜥腳形亞目的研究基礎上，提出一個演化樹，認為原始的農神龍、槽齒龍、埃弗拉士龍代表基礎的外類群（基本上是屬於並系群的槽齒龍科），早於原蜥腳下目與蜥腳下目的分化。在2004年，耶茨提出近蜥龍是最原始的蜥腳類恐龍，儘管牠們的體型為典型的原蜥腳類。但是，近蜥龍的亞成年標本的顱骨特徵，可能會推翻上述理論。黑丘龍科與貝里肯龍科是蜥腳下目演化支中非常原始的物種。

大椎龍的頭顱骨
## 科學檢驗

原蜥腳類的頭顱骨長度接近股骨長度的一半；牠們的頜部關節稍微低於上頜骨的齒列水平線。牠們的牙齒是同型齒或稍微同型齒，牙齒小而呈匙形、邊緣呈粗鋸齒狀。前掌第一指有彎曲的指骨，以及大型、銳利的爪；第二與第三指長度幾乎相等，指骨小型、微彎，帶有指爪；第四與第五指都縮小，而且缺乏指爪。典型原蜥腳類的指骨數量是2-3-4-3。恥骨的末端部分，形成寬廣、平坦的平板。腳部的第五個趾骨是退化的。股骨接近外髁（lateral condyle）側有直條骨脊。股骨粗隆部（Trochanter）的側前面有個由內向外的微弱波紋，而粗隆部的主要部份低於股骨頭。

## 演化歷史

板龍的骨架模型，位於德國圖賓根大學博物館
蜥腳形亞目首次出現於盤古大陸時，是群小型（身長1.5到3公尺）、原始的物種，牠們首次出現於卡尼階中或晚期（晚三疊紀的第一個時期）。牠們發現於巴西（農神龍與黑水龍）、馬達加斯加（最近發現的未命名屬）、以及摩洛哥（愛珍多龍）。

原蜥腳類保持一樣的身體構造，但在諾利階的中期，牠們身體長度已成長到兩倍；例如在德國Stubensandstein地區的中下層發現的纖細板龍（*Plateosaurus gracilis*）身長4到6公尺。纖細板龍之後演化出板龍屬的其他種；板龍身長8公尺，體重約1.5公噸以上，並且是該環境的優勢植食性動物，從諾利階晚期直到瑞提階。同一期間，生存於阿根廷的里奧哈龍，是較大型的原蜥腳類恐龍，也佔據相同的生態位。里奧哈龍身長10公尺，因體型過大而必須以四足行走。同一時代的非洲南部，巨型植食性動物的生態位並非屬於原蜥腳類恐龍，而是基礎蜥腳類恐龍，例如：優肢龍、黑丘龍、貝里肯龍、以及雷前龍。有趣的是，在諾利階到瑞提階這段期間，蜥腳形亞目恐龍佔據者大型植食性動物的生態位，但大型肉食性動物的生態位仍由伪鳄类（鳥鱷科、副鳄形类）佔據。

三疊紀-侏儸紀滅絕事件將槽齒龍科、黑丘龍科、貝里肯龍科、以及里奧哈龍科滅亡了；但近蜥龍科、板龍科、以及真正的蜥腳下目恐龍持續存活者，後兩個生物群演化出更多物種。在侏儸紀早期，蜥腳下目恐龍開始多樣化發展，例如火山齒龍科；原蜥腳下目恐龍也輻射演化出許多中等體型（身長4到6公尺）的植食性物種，例如：大椎龍、祿豐龍、以及雲南龍，並如同牠們的晚三疊紀祖先一樣成功。

原蜥腳下目恐龍的優勢在早侏羅紀的後期結束。雖然至少有三個屬繼續生存至侏羅紀中期（砂龍、祿豐龍、雲南龍），但已并非優勢植食性動物，而蜥腳類恐龍繼續演化出巨大的體型（尤其是真蜥腳類）。

## 分類

濫食龍的骨骼想像圖
在2009年發現的地爪龍，被視為是蜥腳形亞目從二足恐龍演化至四足恐龍的關鍵物種
以下選自於亞當·耶茨（Adam Yates）的2003年研究、以及彼得·加爾東（Peter M. Galton）的2001年研究
[1]（页面存档备份，存于互联网档案馆）：

* **蜥腳形亞目 Sauropodomorpha**
	+ ?愛珍多龍 *Azendohsaurus*
	+ 地爪龍 *Aardonyx*
	+ 農神龍 *Saturnalia*
	+ 槽齒龍 *Thecodontosaurus*
	+ 埃弗拉士龍 *Efraasia*
	+ 濫食龍 *Panphagia*
	+ **原蜥腳下目 PROSAUROPODA**
		- ?易門龍 *Yimenosaurus*
		- ?鼠龍 *Mussaurus*
		- **里奧哈龍科 Riojasauridae 優脛龍 *Eucnemesaurus*
			* 里奧哈龍 *Riojasaurus*
		- **板龍類 Plateosauria **板龍科 Plateosauridae**
				+ ?沙怪龍
				+ 板龍 *Plateosaurus*
				+ 鞍龍 *Sellosaurus*
				+ 黑水龍 *Unaysaurus*
			* **大椎龍科 Massospondylidae**
				+ ?遠食龍 *Adeopapposaurus*
				+ ?冰河龍 *Glacialisaurus*
				+ 科羅拉多斯龍 *Coloradisaurus*
				+ 祿豐龍 *Lufengosaurus*
				+ 大椎龍 *Massospondylus*
				+ 雲南龍 *Yunnanosaurus*
				+ 金山龍 *Jingshanosaurus*

## 種系發生學

在蜥腳形亞目中，原蜥腳下目與蜥腳下目的分類關係、演化位置，隨者兩者的定義與範圍，而產生不同結果。

**耶茨/基欽的2003年版本**

以下演化樹取自於亞當·耶茨（Adam Yates）與詹姆斯·基欽（James Kitching）的2003年研究：

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| 蜥腳形亞目
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|  |  |
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|  | *農神龍*  |
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|  | *槽齒龍*  |
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| --- | --- |
|  | *埃弗拉士龍*  |
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| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| **原蜥腳下目** |

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|  | *里奧哈龍*  |
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| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| 板龍類
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|  | *板龍*  |
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| 大椎龍科
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| --- | --- |
|  | *科羅拉多斯龍* |
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| --- | --- |
|  | *大椎龍*  |
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|  | *祿豐龍*  |
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| 蜥腳下目
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| --- | --- |
|  | *近蜥龍*  |
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| --- | --- |
|  | *黑丘龍* |
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|  | *雷前龍*  |
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|  | *伊森龍* |
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|  | *哥打龍* |
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| unnamed |

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|  | *火山齒龍*  |
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真蜥腳類  |
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**加爾東/阿普徹奇的2004年版本**

以下演化樹取自於彼得·加爾東（Peter M. Galton）與保羅·阿普徹奇（Paul Upchurch）的2004年研究：

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| 蜥腳形亞目
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蜥腳下目  |
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| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| **原蜥腳下目** |

|  |  |
| --- | --- |
|  | *槽齒龍*  |
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| --- | --- |
|  | *農神龍*  |
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| 近蜥龍類
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| --- | --- | --- | --- | --- | --- | --- | --- |
| 近蜥龍科
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|  | *近蜥龍*  |
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|  | *砂龍* |
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| 黑丘龍科
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|  |  |
| --- | --- |
|  | *里奧哈龍*  |
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| --- | --- |
|  | *黑丘龍* |
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|  | *卡米洛特龍*  |
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|  | *萊森龍* |
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| 板龍類
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|  | *金山龍*  |
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|  | *易門龍* |
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|  | *大椎龍*  |
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| 板龍科
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|  |  |
| --- | --- |
|  | *鼠龍*  |
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| --- | --- |
|  | *科羅拉多斯龍* |
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|  | *祿豐龍*  |
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|  | *中國兀龍*  |
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|  | *優肢龍* |
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|  | *鞍龍*  |
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|  | *板龍*  |
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**耶茨的2007年版本**
以下演化樹，取自於亞當·耶茨的2007年研究：

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| 蜥腳形亞目
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|  |  |
| --- | --- |
|  | *農神龍*  |
|  |
| unnamed |

|  |  |
| --- | --- |
|  | *槽齒龍*  |
|  |
|  | *泉水谷龍*  |
|  |
| unnamed |

|  |  |
| --- | --- |
|  | *埃弗拉士龍*  |
|  |
| unnamed |

|  |  |
| --- | --- |
|  | *祖父板龍* |
|  |
| unnamed |

|  |  |
| --- | --- |
|  | *呂勒龍* |
|  |
|  |

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| 板龍類
 |

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| 板龍科
 |

|  |  |
| --- | --- |
|  | *黑水龍*  |
|  |
| unnamed |

|  |  |
| --- | --- |
|  | *纖細板龍*（*鞍龍*）  |
|  |
| unnamed |

|  |  |
| --- | --- |
|  | *恩氏板龍*  |
|  |
|  | *長頭板龍*  |
|  |

 |
|  |

 |
|  |

 |
|  |
| 大腳類
 |

|  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- |
| 里奧哈龍科
 |

|  |  |
| --- | --- |
|  | *里奧哈龍*  |
|  |
|  | *優脛龍* |
|  |

 |
|  |
| unnamed |

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| 大椎龍科
 |

|  |  |
| --- | --- |
|  | *大椎龍*  |
|  |
| unnamed |

|  |  |
| --- | --- |
|  | *科羅拉多斯龍* |
|  |
|  | *祿豐龍*  |
|  |

 |
|  |

 |
|  |
| unnamed |

|  |  |
| --- | --- |
|  | *金山龍*  |
|  |
|  |

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| 近蜥龍類
 |

|  |  |
| --- | --- |
|  | *近蜥龍*  |
|  |
| unnamed |

|  |  |
| --- | --- |
|  | *雲南龍*  |
|  |
| unnamed |

|  |  |
| --- | --- |
|  | *黑丘龍* |
|  |
|  |
蜥腳下目  |
|  |

 |
|  |

 |
|  |

 |
|  |

 |
|  |

 |
|  |

 |
|  |

 |
|  |

 |
|  |

 |
|  |

 |
|  |

 |
|  |

 |
|  |

 |
|  |

 |
|  |

## 參考資料

* Gauffre F.-X. (1993): The Prosauropod Dinosaur Azendohsaurus laaroussii from the Upper Triassic of Morocco. Palaeontology 36(4): 897–908.
* Upchurch, P (1998), The phylogenetic relationships of sauropod dinosaurs. Zool. J. Linnean Soc. 124: 43–103.
* Yates, A. M. (2004) Anchisaurus polyzelus (Hitchcock): the smallest known sauropod dinosaur and the evolution of gigantism among sauropodomorph dinosaurs: Postilla, n. 230, 58 pp.
* Yates, A.M. & Kitching, J. W. (2003) The earliest known sauropod dinosaur and the first steps towards sauropod locomotion. Proc. R. Soc. Lond.: B Biol Sci. 2003 Aug 22; 270(1525): 1753–8.
* Lu, J., Li, T., Zhong, S., Azuma, Y., Fujita, M., Dong, Z., and Ji, Q. (2007). New yunnanosaurid dinosaur (Dinosauria, Prosauropoda) from the Middle Jurassic Zhanghe Formation of Yuanmou, Yunnan Province of China. *Memoir of the Fukui Prefectural Dinosaur Museum* **6**: 1-15.
* Weishampel, D.B., Dodson, P., Osmólska, H. (eds.) (2004). *The Dinosauria, Second Edition*. University of California Press., 861 pp.
* Fedak, T. J. and Galton, P.M. (2007). New information on the braincase and skull of *Anchisaurus polyzelus* (Lower Jurassic, Connecticut, USA; Saurischia: Sauropodomorpha): Implications for sauropodomorph systematics. Special Papers in Palaeontology (77 ) 245-260.

* 原蜥腳下目 Palaeos網站
* 蜥腳形亞目 種系發生學（页面存档备份，存于互联网档案馆） by Mickey Mortimer(2003) - Dinosaur Mailing List Archives論壇