早白垩世热河生物群手盗龙类和原始鸟类的飞羽形态研究

The evolution of feather has been a hard issues in paleontology and evolutionary biology due to the lack of the fossil record. In recent years, the related research has greatly improved the understanding of this issues, and a series of great papers has been published. Nevertheless, there are still many unresolved problems on the study of early feathers. Here in this project, we plan to carefully study the morphology of the feathers and reconstruct their distribution pattern in various taxa including non-avian maniraptorans, such as Caudipteryx, Microraptor and basal bird, Jeholornis, Confuciusornis, Sapeornis as well as early ornithothoraces，Protoptyrex, Archaeorhynchus, Yanornis etc., based on the numerous specimens of feathered dinosaurs and early birds housed in Shandong Tianyu musuem. we will also extensively collect the remiges information of mordern bird, and give close morphology comparation between the mordern birds and fossil ones, find the differences and similarities of these remiges. According to the result of the above comparation, we will analysis the distribution pattern of the main remiges morphotypes in the phylogenetic tree, and try to find the evolutionary rules of remiges. This study can not only provide numerous new morphology information of remiges of many known important taxa including non-avian maniraptorans and early birds from Jehol Biota, and reconstruct the distribution pattern among these taxa, but also increase ouher morphotypes recognized in basal birds and non-avian dinosaurs, and make detail comparison among these morphotypes based on the information of new specimens. we will give a close morphology analysis of all these currently available data and build on current understanding of modern feather cellular and molecular morphogenesis to explore the potential structural mechanism that lead to the extinct of these feather morphotypes. This study will involve into many important Mesozoic taxa discovered in Jehol Group, including the non-avian maniraptorans, such as Microraptor and basal bird, Jeholornis, Confuciusornis, Sapeornis as well as early ornithothoraces，Protoptyrex, Archaeorhynchus, Yanornis etc..This study can provide numerous new information of feather and skeleton morphology by abundant new well-preserved specimens of known important taxa of the dinosaurs and early birds, can not only help to recognize the feathering evolution of the Jehol dinosaurs and birds, but also improve the understanding of the origin and early evolution of birds and flight.

羽毛演化是古生物学和演化生物学的一个研究难题，主要原因在于相关化石记录贫乏。近年来的相关研究极大推动了这一研究方向的进展，并产生了一批具有世界性影响的成果。但是，有关早期飞羽的形态学研究方面还存在许多不足。本项目拟以山东天宇自然博物馆收藏的尾羽龙、小盗龙等非鸟手盗龙类恐龙和热河鸟、孔子鸟、会鸟、原羽鸟、燕鸟等原始鸟类为研究对象，对保存的飞羽进行细致的形态学研究，并精确复原这些属种飞羽的形态特征和分布情况。本研究还将采集现生鸟类飞羽形态信息，并与化石飞羽进行对比，在对比化石和现代飞羽形态相似性和不同点的基础上，分析飞羽主要形态特征在系统演化树上的分布情况，总结飞羽演化规律。本研究不仅能够为人们提供有关热河群的若干重要非鸟手盗龙类和原始鸟类属种飞羽的大量形态信息，复原这些重要属种的飞羽分布情况，而且更有助于人们深入了解这些重要属种的形态和演化，并最终促进人们对于鸟类起源和飞行演化的理解。

羽毛是最复杂的皮肤衍生物，它的起源和演化是一个非常重要但存在高度争议的问题。本项目对天宇自然博物馆馆藏的100多件非鸟手盗龙类和原始基干鸟类进行详细观测和测量，筛选出30多件符合研究条件的标本，在筛选过程中，发现了近10件新物种，已经发表的有4种，包括奇翼龙、无牙古食种鸟、弥曼始今鸟、重明鸟。在近鸟龙、奇翼龙、始孔子鸟等上研究了羽毛形态、翅膀形态以及翅膀前后翼膜等的特点，同时本项目在组织、亚细胞结构、蛋白等大分子结构及元素分析等多方面，进行了现代古生物学的研究尝试，取得了较好的成绩。研究结果多次发表在NATURE、NATURE COMMUNIACTIONS和PNAS等杂志上。. 主要进展有：（1）发现擅攀龙类的奇翼龙(Yi Qi)具有独特的上肢结构、翼膜和羽毛。其羽毛呈丝状，接近原始羽毛，在手指附近发现了残缺翼膜。表明奇翼龙的翅膀主要由翼膜构成，而不是像鸟类及其近亲那样主要由羽毛构成。.（2）通过对一古喙鸟的研究，发现了其尾羽结构与现代鸟类相似，保存有12枚尾羽。同时经过多次试验并使用各种先进仪器设备，克服多重困难，终于首次使早期动物的肺部结构得以较好地呈现。.（3）主要应用激光诱导荧光成像技术，对近鸟龙的肌肉等软组织进行了重建，为以后采用现代工程学的方法对脊椎动物进行“结构-功能”的定量分析打下了重要的基础。.（4）发现最早的始孔子鸟具有翼膜结构，保存有最早记录的小翼羽，也发现了羽毛印痕中保存了黑色素和角蛋白的分子学证据，为化石中很难保存的组织结构的分子保存和分析提供了新的思路。.（5）弥曼始今鸟（Archaeornithura meemannae）的发现。弥曼始今鸟的羽毛保存完整，包括初级和次级飞羽、扇状尾羽，以及在低速飞行时具有重要控制作用的小翼羽。具有较为宽大的羽轴，羽轴中部发育具有纵向凹痕，而其覆羽不具羽轴，表明现生鸟类的上述羽毛特征是在始祖鸟和较进步的今鸟型类中平行演化而来的。