孔子鸟目(鸟类:尾综骨类)的分类厘定（英文）
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摘要
孔子鸟目是早白垩世鸟类中具有角质喙和尾综骨的最原始的鸟类类群之一。在过去近20年的研究历史中,千余件孔子鸟类标本被发现,其数目多过其他中生代鸟类和非鸟类恐龙。目前,共计4个属,10个种被先后归入到孔子鸟目中,然而这些属种的分类位置是否有效尚不清楚。通过对已发表的所有孔子鸟类属种进行详细的对比研究,证实了其中多数属种的鉴定特征有误,这些特征或受保存的影响,或代表了种内和个体发育的变化,亦或广泛分布在孔子鸟类或基干鸟类中,而并不能成为这些属种建属或建种的依据。认为此前命名的孙氏孔子鸟、费氏孔子鸟、义县锦州鸟、张吉营锦州鸟和建昌孔子鸟均属于圣贤孔子鸟的同物异名;而川州孔子鸟未保存确切的鉴定特征而难以和其他孔子鸟类相区别,暂将其归入孔子鸟科未定种。至此,对所有已报道的孔子鸟目的分类厘定认为,目前孔子鸟目共包括1科、3属、4种,分别是圣贤孔子鸟、杜氏孔子鸟、横道子长城鸟和郑氏始孔子鸟,并对这些属种各自的鉴定特征进行了修正。
The Confuciusornithiformes is a basal clade of Early Cretaceous birds that includes the oldest and most basal birds with a toothless beak and an abbreviated bony tail. Over the last two decades, thousands of specimens have been collected, more than for any other group of Mesozoic birds or non-avian dinosaurs. Ten species separated into four genera have been erected with limited taxonomic phylogenetic scrutiny. Here, we perform a comparative study of these ten species, and demonstrate that most of these taxa were originally diagnosed by characters that prove to be either preservational artifacts, intraspecific variations, subject to ontogenetic variation, or widely distributed among the Confuciusornithiformes or a more phylogenetically inclusive group. Our results suggest that ‘Confuciusornis suniae', ‘C. feducciai', ‘Jinzhouornis yixianensis', ‘J. zhangjiyingia', and ‘C. jianchangensis' are all junior synonyms of C. sanctus. ‘C. chuonzhous' lacks autapomorphies of C. sanctus and is referred to Confuciusornithiformes incertae sedis. Our taxonomic reappraisal of published materials indicates that the Confuciusornithiformes consists of one family, three genera, and four species: C. sanctus, C. dui, Changchengornis hengdaoziensis, and Eoconfuciusornis zhengi, for which we provide revised diagnoses.
The Confuciusornithiformes is a basal clade of Early Cretaceous birds that includes the oldest and most basal birds with a toothless beak and an abbreviated bony tail. Over the last two decades, thousands of specimens have been collected, more than for any other group of Mesozoic birds or non-avian dinosaurs. Ten species separated into four genera have been erected with limited taxonomic phylogenetic scrutiny. Here, we perform a comparative study of these ten species, and demonstrate that most of these taxa were originally diagnosed by characters that prove to be either preservational artifacts, intraspecific variations, subject to ontogenetic variation, or widely distributed among the Confuciusornithiformes or a more phylogenetically inclusive group. Our results suggest that ‘Confuciusornis suniae', ‘C. feducciai', ‘Jinzhouornis yixianensis', ‘J. zhangjiyingia', and ‘C. jianchangensis' are all junior synonyms of C. sanctus. ‘C. chuonzhous' lacks autapomorphies of C. sanctus and is referred to Confuciusornithiformes incertae sedis. Our taxonomic reappraisal of published materials indicates that the Confuciusornithiformes consists of one family, three genera, and four species: C. sanctus, C. dui, Changchengornis hengdaoziensis, and Eoconfuciusornis zhengi, for which we provide revised diagnoses.
引文
Agnolín F L,Egli F B,Chatterjee S et al.,2017.Vegaviidae,a new clade of southern diving birds that survived the K/Tboundary.Sci Nat,104:87
Chiappe L M,2002.Basal birds phylogeny:problems and solution.In:Chiappe L M,Witmer L M eds.Mesozoic Birds:Above the Heads of Dinosaurs.Berkeley:University of California Press.448-472
Chiappe L M,Meng Q J,2016.Birds of Stone:Chinese Avian Fossils from the Age of Dinosaurs.Baltimore:Johns Hopkins University Press.1-294
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
Chiappe L M,Ji S A,Ji Q et al.,1999.Anatomy and systematics of the Confuciusornithidae(Theropoda:Aves)from the Late Mesozoic of northeastern China.Bull Am Mus Nat Hist,242:1-89
Chiappe L M,Marugán-Lobón J,Ji S A et al.,2008.Life history of a basal bird:morphometrics of the Early Cretaceous Confuciusornis.Biol Lett,4:719-723
De Ricqlès A J,Padian K,Horner J R et al.,2003.Osteohistology of Confuciusornis sanctus(Theropoda:Aves).J Vert Paleont,23:373-386
Gao C L,Chiappe L M,Zhang F J et al.,2012.A subadult specimen of the Early Cretaceous bird Sapeornis chaoyangensis and a taxonomic reassessment of sapeornithids.J Vert Paleont,32:1103-1112
Gao K Q,Li Q G,Wei M R et al.,2009.Early Cretaceous birds and pterosaurs from the Sinuiju Series,and geographic extension of the Jehol Biota into the Korean Peninsula.J Paleontol Soc Korea,25:57-61
Gill F B,2007.Ornithology,3rd ed.New York:W.H.Freeman.1-720
Goloboff P A,Catalano S A,2016.TNT version 1.5,including a full implementation of phylogenetic morphometrics.Cladistics,32:221-238
He H Y,Wang X L,Zhou Z H et al.,2004.Timing of the Jiufotang Formation(Jehol Group)in Liaoning,northeastern China,and its implications.Geophy Res Lett,31:L12605
He H Y,Wang X L,Jin F et al.,2006.The 40Ar/39Ar dating of the early Jehol Biota from Fengning,Hebei Province,northern China.Geochem Geophy Geosyst,7:Q04001
Hou L H,1997.Mesozoic Birds of China.Nantou Country:Feng-Huang-Ku Bird Park of Taiwan.1-228
Hou L H,Zhou Z H,Gu Y C et al.,1995.Confuciusornis sanctus,a new Late Jurassic sauriurine bird from China.Chin Sci Bull,40:1545-1551
Hou L H,Martin L D,Zhou Z H et al.,1999.A diapsid skull in a new species of the primitive bird Confuciusornis.Nature,399:679-682
Hou L H,Zhou Z H,Zhang F C et al.,2002.Mesozoic Birds from Western Liaoning in China.Shenyang:Liaoning Science and Technology Publishing House.1-120
Hu H,O’Connor J K,2017.First species of Enantiornithes from Sihedang elucidates skeletal development in Early Cretaceous enantiornithines.J Syst Palaeontol,15:909-926
Li D S,2010.Osteology of Confuciusornis sanctus.MS thesis.Beijing:Graduate School of Chinese Academy of Sciences.1-77
Li L,Wang J Q,Hou S L,2010.A new species of Confuciusornis from Lower Cretaceous of Jianchang,Liaoning,China.Glob Geol,29:183-187
Linnaeus C,1758.Systema Naturae per Regna Tria Naturae,Secundum Classes,Ordines,Genera,Species,cum Characteribus,Differentiis,Synonymis,Locis.Vol.1:Regnum Animale.Editio decima,reformata.Stockholm:Laurentii Salvii.1-824
Marugán-Lobón J,Chiappe L M,Ji S A et al.,2011.Quantitative patterns of morphological variation in the appendicular skeleton of the Early Cretaceous bird Confuciusornis.J Syst Palaeontol,9:91-101
Maxwell E E,2008.Comparative embryonic development of the skeleton of the domestic turkey(Meleagris gallopavo)and other galliform birds.Zoology,111:242-257
Navalón G,Meng Q J,Marugán-Lobón J et al.,2018.Diversity and evolution of the Confuciusornithidae:evidence from a new 131-million-year-old specimen from the Huajiying Formation in NE China.J Asian Earth Sci,152:12-22
O’Connor J K,Chiappe L M,2011.A revision of enantiornithine(Aves:Ornithothoraces)skull morphology.J Syst Palaeontol,9:135-157
O’Connor J K,Chiappe L M,Bell A,2011.Pre-modern birds:avian divergences in the Mesozoic.In:Gareth J D,Gary Keds.Living Dinosaurs:the Evolutionary History of Birds.Oxford:Wiley.39-114
O’Connor J K,Wang M,Zheng X T et al.,2014.The histology of two female Early Cretaceous birds.Vert PalAsiat,52:112-128
O’Connor J K,Zheng X T,Sullivan C et al.,2015.Evolution and functional significance of derived sternal ossification patterns in ornithothoracine birds.J Evol Biol,28:1550-1567
Pan Y H,Zheng W J,Moyer A E et al.,2016.Molecular evidence of keratin and melanosomes in feathers of the Early Cretaceous bird Eoconfuciusornis.Proc Natl Acad Sci USA,113:E7900-E7907
Sullivan C,Xu X,2017.Morphological diversity and evolution of the jugal in dinosaurs.Anat Rec,300:30-48
Wang M,Hu H,2017.A comparative morphological study of the jugal and quadratojugal in early birds and their dinosaurian relatives.Anat Rec,300:62-75
Wang M,Zhou Z H,2017a.The evolution of birds with implications from new fossil evidences.In:Maina J N ed.The Biology of the Avian Respiratory System:Evolution,Development,Structure and Function.Cham:Springer International Publishing.1-26
Wang M,Zhou Z H,2017b.A new adult specimen of the basalmost ornithuromorph bird Archaeorhynchus spathula(Aves:Ornithuromorpha)and its implications for early avian ontogeny.J Syst Palaeontol,15:1-18
Wang M,Zhou Z H,O’Connor J K et al.,2014.A new diverse enantiornithine family(Bohaiornithidae fam.nov.)from the Lower Cretaceous of China with information from two new species.Vert PalAsiat,52:31-76
Wang M,O’Connor J K,Pan Y H et al.,2017a.A bizarre Early Cretaceous enantiornithine bird with unique crural feathers and an ornithuromorph plough-shaped pygostyle.Nat Commun,8:14141
Wang M,Li Z H,Zhou Z H,2017b.Insight into the growth pattern and bone fusion of basal birds from an Early Cretaceous enantiornithine bird.Proc Natl Acad Sci USA,114:11470-11475
Wang Y,Hu H,O’Connor J K et al.,2017.A previously undescribed specimen reveals new information on the dentition of Sapeornis chaoyangensis.Cretaceous Res,74:1-10
Zhang F C,Hou L H,Ou Y L,1998.Osteological microstructure of Confuciusornis:preliminary report.Vert PalAsiat,36:126-135
Zhang F C,Zhou Z H,Benton M J,2008.A primitive confuciusornithid bird from China and its implications for early avian flight.Sci China Ser A,51:625-639
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
Zhang Z H,Gao C L,Meng Q J et al.,2009.Diversification in an Early Cretaceous avian genus:evidence from a new species of Confuciusornis from China.J Ornithol,150:783-790
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,3:1116
Zheng X T,O’Connor J,Huchzermeyer F et al.,2013.Preservation of ovarian follicles reveals early evolution of avian reproductive behaviour.Nature,495:507-511
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,4:441-452
Zhou S,Zhou Z H,O’Connor J K,2013.Anatomy of the basal ornithuromorph bird Archaeorhynchus spathula from the Early Cretaceous of Liaoning,China.J Vert Paleont,33:141-152
Zhou Z H,2014.The Jehol Biota,an Early Cretaceous terrestrial Lagerst?tte:new discoveries and implications.Natl Sci Rev,1:543-559
Zhou Z H,Zhang F C,2002.A long-tailed,seed-eating bird from the Early Cretaceous of China.Nature,418:405-409
Zhou Z H,Zhang F C,2003.Anatomy of the primitive bird Sapeornis chaoyangensis from the Early Cretaceous of Liaoning,China.Can J Earth Sci,40:731-747
Zhou Z H,Zhang F C,2006a.Mesozoic birds of China:a synoptic review.Vert PalAsiat,44:74-98
Zhou Z H,Zhang F C,2006b.A beaked basal ornithurine bird(Aves,Ornithurae)from the Lower Cretaceous of China.Zool Scr,35:363-373
Zhou Z H,Zhang F C,Li Z H,2010.A new Lower Cretaceous bird from China and tooth reduction in early avian evolution.Proc R Soc B,277:219-227
Chiappe L M,2002.Basal birds phylogeny:problems and solution.In:Chiappe L M,Witmer L M eds.Mesozoic Birds:Above the Heads of Dinosaurs.Berkeley:University of California Press.448-472
Chiappe L M,Meng Q J,2016.Birds of Stone:Chinese Avian Fossils from the Age of Dinosaurs.Baltimore:Johns Hopkins University Press.1-294
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
Chiappe L M,Ji S A,Ji Q et al.,1999.Anatomy and systematics of the Confuciusornithidae(Theropoda:Aves)from the Late Mesozoic of northeastern China.Bull Am Mus Nat Hist,242:1-89
Chiappe L M,Marugán-Lobón J,Ji S A et al.,2008.Life history of a basal bird:morphometrics of the Early Cretaceous Confuciusornis.Biol Lett,4:719-723
De Ricqlès A J,Padian K,Horner J R et al.,2003.Osteohistology of Confuciusornis sanctus(Theropoda:Aves).J Vert Paleont,23:373-386
Gao C L,Chiappe L M,Zhang F J et al.,2012.A subadult specimen of the Early Cretaceous bird Sapeornis chaoyangensis and a taxonomic reassessment of sapeornithids.J Vert Paleont,32:1103-1112
Gao K Q,Li Q G,Wei M R et al.,2009.Early Cretaceous birds and pterosaurs from the Sinuiju Series,and geographic extension of the Jehol Biota into the Korean Peninsula.J Paleontol Soc Korea,25:57-61
Gill F B,2007.Ornithology,3rd ed.New York:W.H.Freeman.1-720
Goloboff P A,Catalano S A,2016.TNT version 1.5,including a full implementation of phylogenetic morphometrics.Cladistics,32:221-238
He H Y,Wang X L,Zhou Z H et al.,2004.Timing of the Jiufotang Formation(Jehol Group)in Liaoning,northeastern China,and its implications.Geophy Res Lett,31:L12605
He H Y,Wang X L,Jin F et al.,2006.The 40Ar/39Ar dating of the early Jehol Biota from Fengning,Hebei Province,northern China.Geochem Geophy Geosyst,7:Q04001
Hou L H,1997.Mesozoic Birds of China.Nantou Country:Feng-Huang-Ku Bird Park of Taiwan.1-228
Hou L H,Zhou Z H,Gu Y C et al.,1995.Confuciusornis sanctus,a new Late Jurassic sauriurine bird from China.Chin Sci Bull,40:1545-1551
Hou L H,Martin L D,Zhou Z H et al.,1999.A diapsid skull in a new species of the primitive bird Confuciusornis.Nature,399:679-682
Hou L H,Zhou Z H,Zhang F C et al.,2002.Mesozoic Birds from Western Liaoning in China.Shenyang:Liaoning Science and Technology Publishing House.1-120
Hu H,O’Connor J K,2017.First species of Enantiornithes from Sihedang elucidates skeletal development in Early Cretaceous enantiornithines.J Syst Palaeontol,15:909-926
Li D S,2010.Osteology of Confuciusornis sanctus.MS thesis.Beijing:Graduate School of Chinese Academy of Sciences.1-77
Li L,Wang J Q,Hou S L,2010.A new species of Confuciusornis from Lower Cretaceous of Jianchang,Liaoning,China.Glob Geol,29:183-187
Linnaeus C,1758.Systema Naturae per Regna Tria Naturae,Secundum Classes,Ordines,Genera,Species,cum Characteribus,Differentiis,Synonymis,Locis.Vol.1:Regnum Animale.Editio decima,reformata.Stockholm:Laurentii Salvii.1-824
Marugán-Lobón J,Chiappe L M,Ji S A et al.,2011.Quantitative patterns of morphological variation in the appendicular skeleton of the Early Cretaceous bird Confuciusornis.J Syst Palaeontol,9:91-101
Maxwell E E,2008.Comparative embryonic development of the skeleton of the domestic turkey(Meleagris gallopavo)and other galliform birds.Zoology,111:242-257
Navalón G,Meng Q J,Marugán-Lobón J et al.,2018.Diversity and evolution of the Confuciusornithidae:evidence from a new 131-million-year-old specimen from the Huajiying Formation in NE China.J Asian Earth Sci,152:12-22
O’Connor J K,Chiappe L M,2011.A revision of enantiornithine(Aves:Ornithothoraces)skull morphology.J Syst Palaeontol,9:135-157
O’Connor J K,Chiappe L M,Bell A,2011.Pre-modern birds:avian divergences in the Mesozoic.In:Gareth J D,Gary Keds.Living Dinosaurs:the Evolutionary History of Birds.Oxford:Wiley.39-114
O’Connor J K,Wang M,Zheng X T et al.,2014.The histology of two female Early Cretaceous birds.Vert PalAsiat,52:112-128
O’Connor J K,Zheng X T,Sullivan C et al.,2015.Evolution and functional significance of derived sternal ossification patterns in ornithothoracine birds.J Evol Biol,28:1550-1567
Pan Y H,Zheng W J,Moyer A E et al.,2016.Molecular evidence of keratin and melanosomes in feathers of the Early Cretaceous bird Eoconfuciusornis.Proc Natl Acad Sci USA,113:E7900-E7907
Sullivan C,Xu X,2017.Morphological diversity and evolution of the jugal in dinosaurs.Anat Rec,300:30-48
Wang M,Hu H,2017.A comparative morphological study of the jugal and quadratojugal in early birds and their dinosaurian relatives.Anat Rec,300:62-75
Wang M,Zhou Z H,2017a.The evolution of birds with implications from new fossil evidences.In:Maina J N ed.The Biology of the Avian Respiratory System:Evolution,Development,Structure and Function.Cham:Springer International Publishing.1-26
Wang M,Zhou Z H,2017b.A new adult specimen of the basalmost ornithuromorph bird Archaeorhynchus spathula(Aves:Ornithuromorpha)and its implications for early avian ontogeny.J Syst Palaeontol,15:1-18
Wang M,Zhou Z H,O’Connor J K et al.,2014.A new diverse enantiornithine family(Bohaiornithidae fam.nov.)from the Lower Cretaceous of China with information from two new species.Vert PalAsiat,52:31-76
Wang M,O’Connor J K,Pan Y H et al.,2017a.A bizarre Early Cretaceous enantiornithine bird with unique crural feathers and an ornithuromorph plough-shaped pygostyle.Nat Commun,8:14141
Wang M,Li Z H,Zhou Z H,2017b.Insight into the growth pattern and bone fusion of basal birds from an Early Cretaceous enantiornithine bird.Proc Natl Acad Sci USA,114:11470-11475
Wang Y,Hu H,O’Connor J K et al.,2017.A previously undescribed specimen reveals new information on the dentition of Sapeornis chaoyangensis.Cretaceous Res,74:1-10
Zhang F C,Hou L H,Ou Y L,1998.Osteological microstructure of Confuciusornis:preliminary report.Vert PalAsiat,36:126-135
Zhang F C,Zhou Z H,Benton M J,2008.A primitive confuciusornithid bird from China and its implications for early avian flight.Sci China Ser A,51:625-639
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
Zhang Z H,Gao C L,Meng Q J et al.,2009.Diversification in an Early Cretaceous avian genus:evidence from a new species of Confuciusornis from China.J Ornithol,150:783-790
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,3:1116
Zheng X T,O’Connor J,Huchzermeyer F et al.,2013.Preservation of ovarian follicles reveals early evolution of avian reproductive behaviour.Nature,495:507-511
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,4:441-452
Zhou S,Zhou Z H,O’Connor J K,2013.Anatomy of the basal ornithuromorph bird Archaeorhynchus spathula from the Early Cretaceous of Liaoning,China.J Vert Paleont,33:141-152
Zhou Z H,2014.The Jehol Biota,an Early Cretaceous terrestrial Lagerst?tte:new discoveries and implications.Natl Sci Rev,1:543-559
Zhou Z H,Zhang F C,2002.A long-tailed,seed-eating bird from the Early Cretaceous of China.Nature,418:405-409
Zhou Z H,Zhang F C,2003.Anatomy of the primitive bird Sapeornis chaoyangensis from the Early Cretaceous of Liaoning,China.Can J Earth Sci,40:731-747
Zhou Z H,Zhang F C,2006a.Mesozoic birds of China:a synoptic review.Vert PalAsiat,44:74-98
Zhou Z H,Zhang F C,2006b.A beaked basal ornithurine bird(Aves,Ornithurae)from the Lower Cretaceous of China.Zool Scr,35:363-373
Zhou Z H,Zhang F C,Li Z H,2010.A new Lower Cretaceous bird from China and tooth reduction in early avian evolution.Proc R Soc B,277:219-227