未成熟羽毛在早白垩世热河鸟类生物群幼年反鸟类中的首次报道
收稿日期: 2019-04-15
网络出版日期: 2020-01-20
基金资助
国家自然科学基金资助(41688103);国家自然科学基金资助(41372014)
版权
First report of immature feathers in juvenile enantiornithines from the Early Cretaceous Jehol avifauna
Received date: 2019-04-15
Online published: 2020-01-20
Copyright
换羽——将一种羽毛替换成另一种的过程——对鸟类具有非常重要的生物学作用。这一过程能够每年将受损的羽毛替换掉,产生与个体发育相关的羽毛类型,或产生处在繁殖活跃期成年个体所具有的覆羽类型。处于萌发阶段的未成熟羽毛被角质鞘包裹,外形呈狭窄管状,没有明显特征。角质鞘完全脱落标志着羽毛的成熟。虽然热河生物群发现了大量与皮肤衍生物相关的化石,但是确切无疑的未成熟羽毛还未有报道,尽管九佛堂组(120 Ma)发现的一件窃蛋龙类——似尾羽龙,保存了疑似未成熟的羽毛。一件缅甸琥珀(9 Ma)中的反鸟幼崽保存了处于萌发阶段的羽毛,由于是三维保存,对外皮结构的解读更为直观。描述了发现于九佛堂组的4件反鸟类幼年个体上保存的疑似未成熟羽毛。与现生鸟类相似,上述疑似未成熟羽毛的近端窄,外形没有明显结构,仅在末端显示出分叉的羽支。认为此前报道的反鸟类多齿胫羽鸟(Cruralispennia multidonta)掌部和胫跗骨上的相似类型的羽毛可能也是未成熟羽毛。未成熟羽毛和与性双型相关的装饰性羽毛在反鸟类幼年个体上的同时出现,可能说明新鸟类具有的复杂的换羽模式——与性双型相关的装饰性羽毛在个体达到性成熟若干年后(经历了若干次换羽后)才会出现,并且只在更接近鸟类冠群的一些类群中出现。
邹晶梅, 福雅曼, 王敏, 郑晓廷 . 未成熟羽毛在早白垩世热河鸟类生物群幼年反鸟类中的首次报道[J]. 古脊椎动物学报, 2020 , 58(1) : 24 -44 . DOI: 10.19615/j.cnki.1000-3118.190823
Molting—the process replacing one plumage with another—is a critically important biological function in Aves. This process annually replaces the feather coat, damaged by normal wear and tear, produces ontogenetic changes in feathering, and produces alternate breeding plumages associated with reproductive activity in adults. Immature, growing feathers are encased in a keratinous sheath, giving them a narrow, tubular, and featureless appearance. The complete loss of the sheath indicates the feather is mature. Despite the wealth of integumentary data published from the Jehol Biota, immature feathers have never been definitively reported, although they may potentially be preserved in a juvenile specimen of the non-avian oviraptorosaur theropod dinosaur Similicaudipteryx from the 120 Ma Jiufotang Formation. A developing feather has been reported in a 99 Ma enantiornithine neonate preserved in Burmese amber, in which three-dimensional preservation makes interpretations of integumentary structures more straightforward. Here we report on probable immature feathers in four juvenile enantiornithines (Aves: Ornithothoraces) from the Jehol Group. As observed in developing feathers in extant birds, the purported immature fossil feathers appear proximally narrow and featureless with barbs protruding only distally. Based on our observations, we suggest that similar-appearing feather structures preserved on the manus and tibiotarsus in the holotype of the enantiornithine Cruralispennia multidonta may alternatively be interpreted as immature feathers. The presence of immature feathers in combination with sexually dimorphic ornamental feathers in juvenile enantiornithines suggests the complex molting patterns of Neornithes, in which such ornaments only appear after several years (following several molts) when reproductive activity is achieved, are limited to a subset of crownward avians.
Key words: Mesozoic; Jehol Biota; Aves; pin feather; plumage; molt
[1] | Amadon D, 1966. Avian plumages and molts. Condor, 68(3):263-278 |
[2] | Chiappe L M, Walker C A, 2002. Skeletal morphology and systematics of the Cretaceous Euenantiornithes (Ornithothoraces: Enantiornithes). In: Chiappe L M, Witmer L M eds. Mesozoic Birds: Above the Heads of Dinosaurs. Berkeley: University of California Press. 240-267 |
[3] | de Souza Carvalho I, Novas F E, Agnolín F L et al., 2015. A Mesozoic bird from Gondwana preserving feathers. Nat Commun, 6:7141 |
[4] | Elzanowski A, 1981. Embryonic bird skeletons from the Late Cretaceous of Mongolia. Palaeontol Pol, 42:147-179 |
[5] | Foth C, 2012. On the identification of feather structures in stem-line representatives of birds: evidence from fossils and actuopalaeontology. Pal?ontol Z, 86:91-102 |
[6] | Gill F B, 2007. Ornithology, 3rd ed. New York: W. H. Freeman and Company. 1-758 |
[7] | Hu H, O’Connor J K, 2017. First species of Enantiornithes from Sihedang elucidates skeletal development in Early Cretaceous enantiornithines. J Syst Palaeontol, 15(11):909-926 |
[8] | Jones D, G?th A, 2008. Mound-builders. Collingwood, Australia: CSIRO Publishing. 1-119 |
[9] | Kaye T G, Falk A R, Pittman M et al., 2015. Laser-stimulated fluorescence in paleontology. PLoS ONE, 10(5):e0125923 |
[10] | Knoll F, Chiappe L M, Sanchez S et al., 2018. A diminutive perinate European Enantiornithes reveals an asynchronous ossification pattern in early birds. Nat Commun, 9:937 |
[11] | Li Q G, Clarke J A, Gao K Q et al., 2018. Elaborate plumage patterning in a Cretaceous bird. PeerJ, 6:e5831 |
[12] | Lovette I J, Fitzpatrick J W, 2004. The Handbook of Bird Biology. Princeton, New Jersey: Princeton University Press. 1-716 |
[13] | Lucas S G, Stettenheim P R, 1972. Avian Anatomy: Integument Part I. Washington D.C.: U.S. Department of Agriculture. 1-340 |
[14] | Murphy M E, King J R, 1986. Composition and quantity of feather sheaths produced by white-crowned sparrows during the postnuptial molt. The Auk, 103(4):822-825 |
[15] | O’Connor J K, Chang H L, 2015. Hindlimb feathers in paravians: primarily ‘wings’ or ornaments? Biol Bull, 42(7):616-621 |
[16] | O’Connor J K, Chiappe L M, Chuong C M et al., 2012. Homology and potential cellular and molecular mechanisms for the development of unique feather morphologies in early birds. Geosciences, 2:157-177 |
[17] | O’Connor J K, Wang M, Zheng X T et al., 2014. The histology of two female Early Cretaceous birds. Vert PalAsiat, 52(1):112-128 |
[18] | O’Connor J K, Wang X L, Sullivan C et al., 2018. First report of gastroliths in the Early Cretaceous basal bird Jeholornis. Cretaceous Res, 84:200-208 |
[19] | Prum R O, 2010. Moulting tail feathers in a juvenile oviraptorisaur. Nature, 468:E1, doi: 10.1038/nature09480 |
[20] | Starck J M, 1993. Evolution of avian ontogenies. In: Power D M ed. Current Ornithology, Vol. 10. New York: Plenum Press. 275-366 |
[21] | Sullivan C, Xu X, O’Connor J K, 2017. Complexities and novelties in the early evolution of avian flight, as seen in the Mesozoic Yanliao and Jehol Biotas of Northeast China. Palaeoworld, 26(2):212-229 |
[22] | Swank W G, 1955. Feather molt as an ageing technique for mourning doves. J Wildlife Manage, 19(3):412-414 |
[23] | Wang M, O’Connor J K, Pan Y H et al., 2017. A bizarre Early Cretaceous enantiornithine bird with unique crural feathers and an ornithuromorph plough-shaped pygostyle. Nat Commun, 8:1-12 |
[24] | Wang X L, O’Connor J K, Zheng X T et al., 2014. Insights into the evolution of rachis dominated tail feathers from a new basal enantiornithine (Aves: Ornithothoraces). Biol J Linn Soc, 113(3):805-819 |
[25] | Wang X L, O’Connor J K, Maina J N et al., 2018. Archaeorhynchus preserving significant soft tissue including probable fossilized lungs. Proc Natl Acad Sci USA, 115:11555-11560 |
[26] | Xing L D, McKellar R C, Wang M et al., 2016. Mummified precocial bird wings in mid-Cretaceous Burmese amber. Nat Commun, 7:12089 |
[27] | Xing L D, O’Connor J K, McKellar R C et al., 2017. A mid-Cretaceous enantiornithine (Aves) hatchling preserved in Burmese amber with unusual plumage. Gondwana Res, 49:264-277 |
[28] | Xing L D, Cockx P, McKellar R C et al., 2018. Ornamental feathers in Cretaceous Burmese amber: resolving the enigma of rachis-dominated feather structure. J Palaeogeogr, 7(13):1-18 |
[29] | Xu X, Zheng X T, You H L, 2010a. Exceptional dinosaur fossils show ontogenetic development of early feathers. Nature, 464:1339-1341 |
[30] | Xu X, Zheng X T, You H L, 2010b. Reply: moulting tail feathers in a juvenile oviraptorosaur. Nature, 468:E2 |
[31] | Zhang F C, Zhou Z H, Dyke G J, 2006. Feathers and ‘feather-like’ integumentary structures in Liaoning birds and dinosaurs. Geol J, 41:395-404 |
[32] | Zhang F C, Kearns S L, Orr P J et al., 2010. Fossilized melanosomes and the colour of Cretaceous dinosaurs and birds. Nature, 463:1075-1078 |
[33] | Zheng X T, Wang X L, O’Connor J K et al., 2012. Insight into the early evolution of the avian sternum from juvenile enantiornithines. Nat Commun, 3:1-8 |
[34] | Zheng X T, Zhou Z H, Wang X L et al., 2013. Hind wings in basal birds and the evolution of leg feathers. Science, 339:1309-1312 |
[35] | Zheng X T, O’Connor J K, Wang X L et al., 2017. Exceptional preservation of soft tissue in a new specimen of Eoconfuciusornis and its biological implications. Natl Sci Rev, 4(3):441-452 |
[36] | Zhou Z H, Zhang F C, 2004. A precocial avian embryo from the Lower Cretaceous of China. Science, 306:653 |
/
〈 | 〉 |