现生鸟类和孔子鸟(Confuciusornis)叉骨软骨的研究：初步分析以及对中生代鸟类飞行方式的启示

doi:10.19615/j.cnki.1000-3118.201222

摘要/Abstract

摘要：

飞行的早期演化是古生物研究的热点问题之一。现生鸟类演化出多种不同飞行方式,然而对中生代鸟类飞行方式的研究亟需寻找具有指示意义的指标。骨骼与关节组织尤其是乌喙骨-叉骨关节是实现鸟类飞行功能的重要组成,因此其关节的组织形态可能有助于反映现生鸟类的飞行方式。鸟类膜质骨中的次级软骨受到表观遗传的重要影响,只能在关节受到肌肉运动刺激的情况下形成,因此能够反映鸟类关节的形成力学环境;对三种不同飞行方式的现生鸟类(珠颈斑鸠、树麻雀和普通楼燕), 以及热河生物群中最为常见的古鸟类之一孔子鸟(Confuciusornis)的叉骨乌喙骨关节的组织学特征进行了分析,显示在所有三种现生鸟类中,叉骨与乌喙骨之间均存在可动关节,并且在叉骨上存在次级软骨。而孔子鸟的叉骨上也存在次级软骨,这是次级软骨组织在中生代鸟类叉骨化石中的首次报道。进一步分析发现,不同现生鸟类物种的次级软骨组织形态存在差异,还需更多数据建立相关形态功能关系以便用于帮助推断中生代鸟类的飞行方式。

关键词: 飞行演化, 鸟类, 孔子鸟, 叉骨, 组织学, 次级软骨

Abstract:

The early evolution of flight is one of the most studied topics in vertebrate paleontology. Living birds have evolved to utilize a variety of flight styles, but studies focused on inferring flight strategies in Mesozoic birds are often contradictory and without a clear consensus, making it necessary to find additional informative characteristics that can be useful for inferences in fossils. Virtually nothing is known about the histology of the avian pectoral girdle, even though skeletal and joint tissues are key candidates to solve form-function relationships. Avian secondary cartilage found on the dermal bones of the avian skeleton is influenced by epigenetics and only forms when joints are stimulated by muscle contractions. As the only dermal bone in the avian postcranium, the furcula is a potential site for the formation of furcular secondary cartilage and merits further attention. It is still unknown whether adult living birds and fossil birds have furcular secondary cartilage. Here we present histological analyses conducted on the furcula-coracoid articulation in three living birds (Spilopelia chinensis, the Spotted dove; Passer montanus, the Eurasian tree sparrow; and Apus apus, the Common swift), taxa that utilize different flight styles, and one of the most common fossil birds of the Jehol Biota, Confuciusornis. Secondary cartilage was identified on the furculae of the living birds and of Confuciusornis, representing the first report of furcular secondary cartilage in the fossil record. Clear differences in secondary cartilage morphologies were observed in the living species, but additional data is required to establish a strong form-function relationship that could be useful for making inferences in Mesozoic birds.

Key words: Evolution of flight, birds, Confuciusornis, furcula, histology, secondary cartilage

中图分类号:

Q915.865

吴倩, 邹晶梅, 李志恒, 艾莉达. 现生鸟类和孔子鸟(Confuciusornis)叉骨软骨的研究：初步分析以及对中生代鸟类飞行方式的启示. 古脊椎动物学报, 2021, 59(2): 106-124.

WU Qian, Jingmai K. O’CONNOR, LI Zhi-Heng, Alida M. BAILLEUL. Cartilage on the furculae of living birds and the extinct bird Confuciusornis: a preliminary analysis and implications for flight style inferences in Mesozoic birds. Vertebrata Palasiatica, 2021, 59(2): 106-124.

图/表 3

Fig. 1 Histology of the furcula-coracoid joint in the Spotted dove, the Eurasian tree sparrow and the Common swift with paraffin slides stained with a modified Masson’s trichrome A, D, G. images of the joint in the Spotted dove; B, E, H. images of the joint in the Eurasian tree sparrow; C, F, I. images of the joint in the Common swift. A-C. show the synovial joints at low resolution; D-F. close-ups on the primary cartilage on the coracoid; G-I. close-ups on the secondary cartilage on the furcula Abbreviations: CO. coracoid; CSC. calcified secondary cartilage; FT. fibrous tissue; FU. furcula; PC. primary cartilage; SC. secondary cartilage; syc. synovial cavity; sym. synovial membrane

Fig. 2 Morphology, CT-scans and histology of the furcula of Confuciusornis (IVPP V 11521) A. photograph of the furcula; B. CT-scan of the left epicleideal process of the furcula; C. demineralized piece of cartilage from the left epicleideal process after 48 hours in EDTA; D. longitudinal ground-section in the right epicleideal process of the furcula; E-G. close-ups of that epicleideal process showing calcified secondary cartilage; H-I. close-ups on the rami seen in D, shown under natural transmitted light (H) and cross-polarized light (I) Abbreviations: CB. cortical bone; CSC. calcified secondary cartilage; epi. epicleideum; FU. furcula; HU. humerus; ICL. inner circumferential layer; MC. medullary cavity; OCL. outer circumferential layer; SC. secondary cartilage; sed. sediment

Fig. 3 Morphology and histology of the scapulocoracoid of Confuciusornis (IVPP V 11521) A. photograph of the fused scapulocoracoid showing the coracoid on one side (CO), the scapula (SCA) on the other side, and the glenoid fossa (gf) for the articulation of the humerus in between; B. a ground-section (cross section) through the glenoid fossa and on the opposite side, remnants of calcified fibrocartilage (CFC) can be seen with some sediment (sed) in a concave area; C-E. close-ups on the fibrocartilage, a few chondrocyte lacunae can be seen as well (blue arrows)

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