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侏羅紀善攀鳥龍類揭示膜質翅膀在恐龍中的演化

文章來源:古脊椎動物與古人類研究所   發布時間:2019-05-09  【字號:     】  

  2019年《自然》(Nature)以封面文章發表了中國科學院古脊椎動物與古人類研究所王敏、鄒晶梅、徐星、周忠和的研究成果:侏羅紀善攀鳥龍類揭示膜質翅膀在恐龍中的演化,展示了在恐龍—鳥類演化歷程中出現大量意想不到的適應飛行的嘗試,與之對應演化出差異顯著的骨骼—表皮衍生物組合。

  在脊椎動物漫長的演化史中,翼龍、鳥類和蝙蝠獨立演化出了形態迥異的飛行結構。相較翼龍和蝙蝠不完整的化石記錄,隨著不斷發現的帶羽毛恐龍和早期鳥類化石,尤其得益于我國中晚侏羅世的燕遼生物群和早白堊世的熱河生物群,有關鳥類飛行起源這一重要科學問題取得了重要進展,而善攀鳥龍類的發現則揭示了“一條匪夷所思的征服藍天之旅”。善攀鳥龍類(Scansoriopterygidae)是恐龍家族中最為怪異的類群,生活在中—晚侏羅世,迄今發現的僅有三個屬種:寧城樹棲龍(Epidendrosaurus ningchengensis)、胡氏耀龍(Epidexipteryx hui)和奇翼龍(Yi qi)。善攀鳥龍類形態特殊,如頭骨高聳、四肢纖細、第三手指(最外側的手指)加長、古老的2-3-4手指指式、尾骨縮短等,儼然是恐龍和鳥類的“混合體”,而它一度被認為是和鳥類具有最近親緣關系的獸腳類恐龍。但上述標本或不完整,或屬于幼年個體,大量形態特征難以觀察,造成它在演化樹上的位置撲朔迷離。2015年徐星等命名的奇翼龍更加為這一類群增添神秘色彩。奇翼龍的前肢附著翼膜,還具有一根棒狀長骨,這樣的長骨在其它恐龍(包括鳥類)中沒有對應的同源結構。因此,奇翼龍被復原成類似翼龍那樣具有膜質翅膀而能夠滑翔。但奇翼龍的標本僅有一件,保存不完整,因此對于棒狀長骨和翼膜的結構還存有爭議。

  2017年周忠和帶領的基礎科學中心團隊在遼寧晚侏羅世地層考察時獲得一件新化石。經過長達一年的室內修理、實驗和對比研究,研究團隊認為其代表一新的善攀鳥龍類,將其命名為長臂渾元龍(Ambopteryx longibrachium)(意指翼龍那樣膜質翅膀和恐龍的混合體)。渾元龍發現于燕遼生物群晚侏羅世早期的海房溝組(距今約1.63億年),其正型標本是目前已知最完整的善攀鳥龍類化石,提供大量形態和生態學信息。渾元龍體長約32厘米,體重約306克。渾元龍在肱骨近端關節面、手指和腰帶形態方面明顯不同于其它善攀鳥龍類,并且具有原始鳥類那樣的尾綜骨,如此縮短的尾骨能進一步將身體重心前移,有利于在飛行/滑翔時保持穩定。更為重要的是,研究人員在渾元龍上發現了和奇翼龍相似的棒狀長骨和翼膜(翼膜中保存有色素體),這一新發現為棒狀長骨和翼膜在善攀鳥龍類中的出現提供了確切無疑的證據。渾元龍體內保存有胃石和疑似尚未完全消化的骨質胃容物,這是在善攀鳥龍類中首次發現的與食性相關的證據,研究人員推測其為雜食性。

  渾元龍的前肢異常加長,甚至超過了中生代多數鳥類。研究人員在對比恐龍前肢時,發現善攀鳥龍類的前肢比例構成非常奇特,而這樣的差異是否與翼膜的出現有關?為了證實這一猜測,王敏等采用基于系統發育關系的主成分分析方法來討論中生代虛骨龍類(包括鳥類)四肢長度的演化,特別是在接近飛行起源時有哪些顯著變化。系統發育主成分分析是在傳統的主成分分析上剔除親緣關系,最大程度保證采樣點的獨立性,同時復原祖先節點的特征狀態,從而展現不同類群的演化趨勢。研究結果顯示,自副鳥類(Paraves,即包括所有鳥類,但不包括竊蛋龍類的最廣義類群)開始,前肢開始加長,但僅有善攀鳥龍類的加長程度接近中生代鳥類,而這一程度其它非鳥龍類恐龍從未獲得。善攀鳥龍類前肢的加長主要源自肱骨和尺骨;在鳥類、馳龍類或者傷齒龍類中,則是掌骨的加長,而這些類群的前肢具有飛羽。研究人員認為善攀鳥龍類通過加長的肱骨和尺骨、第三手指,與棒狀長骨來附著膜質的翅膀,而鳥類、馳龍類和傷齒龍類則需要較長的掌骨來附著飛羽,顯示出兩種不同的飛行模式(“膜質翅膀和短掌骨”、“羽毛翅膀和長掌骨”)對前肢結構產生的巨大改變。

  已知的善攀鳥龍類均生活在晚侏羅世,類似的膜質翅膀沒有在白堊紀的恐龍中出現。由飛羽構成的翅膀自晚侏羅世出現就延續到白堊紀,進過漫長的演化最終形成了鳥類的羽翼,使后者成為多樣性最豐富的現生四足動物。善攀鳥龍類獨特的飛行結構代表了飛行演化的一次短暫嘗試。

  該研究得到國家自然科學基金基礎科學中心項目(克拉通破壞與陸地生物演化)、優秀青年科學基金項目、中科院青年創新促進會以及巖石圈演化國家重點實驗室(華北克拉通破壞與燕遼-熱河生物群演化)的支持。

 

1. 《自然》當期雜志封面:古脊椎所發現侏羅紀具膜質翅膀的恐龍 

 

  2. a, 長臂渾元龍正型標本;b, 骨骼復原圖;c. 膜質翼膜色素體;d. 骨質胃容物組織切片. bn, 骨質胃容物; gs, 胃石; mw, 膜質翼膜; st, 棒狀長骨(王敏供圖) 

 

  3. 中生代虛骨龍類系統樹和肢骨演化:在飛行起源時,副鳥類不同類群顯示出不同的前肢加長過程:善攀鳥龍類通過肱骨加長,結合第三手指和棒狀長骨附著膜質翅膀;而馳龍類、傷齒龍類和鳥類則通過掌骨的加長以附著羽狀翅膀(王敏供圖)

 

4. 長臂混元龍復原圖(張宗達供圖)



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