孔子鸟目(鸟类：尾综骨类)的分类厘定

doi:10.19615/j.cnki.1000-3118.180530

摘要/Abstract

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。

关键词: 白垩纪, 鸟类, 孔子鸟目, 孔子鸟属, 分类, 系统发育

Abstract:

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.

Key words: Cretaceous, Aves, Confuciusornithiformes, Confuciusornis, taxonomy, phylogeny

中图分类号:

Q915.865

王敏, 邹晶梅, 周忠和. 孔子鸟目(鸟类：尾综骨类)的分类厘定. 古脊椎动物学报, 2019, 57(1): 1-37.

WANG Min, Jingmai O’CONNOR, ZHOU Zhong-He. A taxonomical revision of the Confuciusornithiformes (Aves: Pygostylia). Vertebrata Palasiatica, 2019, 57(1): 1-37.

图/表 15

Fig. 1 Skull of Confuciusornis sanctus in comparison with those of C. dui and other Early Cretaceous birds A, C. photograph (A) and line drawing (C) of Confuciusornis sanctus IVPP V 13168 with a separate drawing of the maxilla (right), showing the halberd-shaped dorsal process of the maxilla (considered an autapomorphy here); B. C. sanctus V 13171 with line drawing of the maxilla; D. drawing of C. dui with a separate drawing of the maxilla (modified from Hou et al., 1999); E. Jeholornis sp. V 14978 (Aves, non-Pygostylia); F. Pterygornis dapingfangensis V 16363 (Aves, Enantiornithes). The arrowheads in B denote the rostral notches of the premaxillary and mandibular symphyses. The arrow in B indicates the ventral process of the surangular (considered an autapomorphy here). Scale bars = 10 mm; C, D not to scale

Fig. 2 Detail anatomy of Confuciusornis sanctus showing some diagnostic characters A. right forelimb of C. sanctus IVPP V 10928 (hand in ventral view, humerus in cranial view); B. hindlimbs of C. sanctus V 11552 in dorsal view; C, D. photograph (C) and reconstruction (D) of the sternum of C. sanctus in dorsal view; E. sternal reconstruction of C. dui (modified from Hou et al., 1999); F. sternal reconstruction of Eoconfuciusornis zhengi (modified from Zhang et al., 2008) The arrowhead in A denotes the ridge on the craniaodorsal margin of the deltopectoral crest, and the arrow indicates the unfused carpometacarpus. The arrowheads in B mark the extensor retinaculum tubercle on the dorsal surface of the distal tibiotarsus (diagnostic character); and the arrows indicate the dorsal tubercles on metatarsals II and III. The arrows in D denote the cranial and caudal projections of the lateral process of the sternum in C. sanctus (diagnostic character). The dash lines in E indicate the length (vertical) and width (horizontal) of the sternum that are measured in this study Scale bars equal 10 mm in A and C, 5 mm in B; D-F not to scale

Fig. 3 Photographs of Confuciusornis sanctus [= 'Confuciusornis chuonzhous'], IVPP V 10919 A. slab; B. counter slab. The broken piece of the proximal end of metatarsal II was misinterpreted as metatarsal V by Hou (1997). See text for further explanations. Scale bars = 10 mm

Fig. 4 Photograph of the holotype of Changchengornis hengdaoziensis, GMV 2129 Scale bars = 20 mm

Fig. 5 Cranial morphology of Changchengornis hengdaoziensis, GMV 2129 A, B. skull in right lateral view preserved on GMV 2129a; C, D. skull in left lateral view preserved on GMV 2129b. A, C. photograph; B, D. line drawing. The dorsal process of the maxilla (labeled as 'mx') identified in Chiappe et al. (1999) is reinterpreted to be the lacrimal here (see text for explanation) Scale bars = 10 mm (A, C); line drawings (C, D) are not to scale

Fig. 6 Pectoral girdle of Changchengornis hengdaoziensis in comparison with Confuciusornis sanctus A. pectoral girdle of Ch. hengdaoziensis in dorsal view; B. line drawing of the furcula (left) and caudal end of the sternum (right) of Ch. hengdaoziensis, showing that the dorsal surface is excavated by a dorsal furrow as in C. sanctus; C. furcula of C. sanctus IVPP V 11521 in dorsal view with drawing of the cross section Scale bars = 10 mm (A, C); line drawings (B, C) are not to scale

Fig. 7 Skeleton and anatomy of Confuciusornis sanctus [= 'Jinzhouornis yixianensis'], IVPP V 14412 A. overview; B. skull in oblique right lateral view; C. proximal left humerus showing the ridge on the cranial surface (arrow), as in C. sanctus; D. lower legs in dorsal view, exhibiting the presences of an extensor retinaculum on distal tibiotarsus (arrowhead), and the proximal ends of metatarsals II and III (arrows), as in C. sanctus. Scale bars = 10 mm

Fig. 8 Skeleton and cranial anatomy of Confuciusornis sanctus [='Jinzhouornis zhangjiyingia'], IVPP V 12352 A. overview; B, C. photograph (B) and line drawing (C) of the skull in left lateral viewThe arrowhead in B denotes the ventral end of the ventral process of the postorbital, which was misinterpreted as the quadratojugal by Hou (1997). Scale bars = 10 mm

Fig. 9 Detail anatomy of Confuciusornis sanctus [= 'Jinzhouornis zhangjiyingia'], IVPP V 12352 A, B. pectoral girdle, sternum, and left forelimb; C, D. hindlimb showing the presence of an extensor retinaculum on distal tibiotarsus (arrow); A, C. photographs; B, D. line drawing. The arrow in A denotes the ridge on the cranial surface of the deltopectoral crest, as in C. sanctus. Scale bars = 10 mm

Fig. 10 Skeletal anatomy of the holotype of Eoconfuciusornis zhengi, IVPP V 11977 A. overview; B, C. photograph (B) and line drawing (C) of the skull; D. furcula (dorsal view) and proximal part of right humerus (cranial view). The arrowhead in C indicates that the surangular bears a ventral process as in C. sanctus. The arrow in C shows that the dorsal process of the maxilla is craniocaudally compressed Scale bars = 10 mm (A, B, D); line drawing (C) is not to scale

Fig. 11 Comparison of furcula and humerus anatomy between Eoconfuciusornis zhengi and Confuciusornis sanctus A. holotype of Eoconfuciusornis zhengi, IVPP V 11977; B. BMNHC-PH870, referred to E. zhengi in Navalón et al. (2018); C. C. sanctus. The arrowheads in B and C indicate that the deltopectoral crest is perforated by a suboval fenestra in BMNHC-PH870 and C. sanctus, but not in E. zhengi. Line drawings are not to scale

Fig. 12 Skeleton and anatomy of Confuciusornis sanctus [= 'Confuciusornis feducciai'], DNHM D2454 A. overview; B. skull in ventral view, showing that the mandibular symphysis is notched rostrally (arrow); C. furcula in ventral view; D. sternum in ventral view, showing the incomplete caudal end (dashed line) and a cleft at the midpoint of the rostral margin (arrow); E. pelvis in ventral viewScale bars equal 20 mm in A, 10 mm in B-E

Fig. 13 Detail anatomy of Confuciusornis sanctus [= 'Confuciusornis feducciai'], DNHM D2454, showing that it is morphologically identical to C. sanctus A. right humerus in cranial view showing the ridge on the proximal part of the cranial surface of the deltopectoral crest (arrow); B. proximal left humerus in cranial view showing the perforated deltopectoral crest; C. dorsal view of metacarpals; D. dorsal view of metatarsals The arrows in D denote the dorsal tubercles on metatarsals II and III. Scale bars = 10 mm

Fig. 14 Skeleton and anatomy of Confuciusornis sanctus [='Confuciusornis jianchangensis'], PMOL-AB00114 A. overview; B. skull; C. thoracic vertebrae showing that the centra are excavated by lateral suboval fossae (arrow), as in C. sanctus; D. hindlimb. The arrows in D denote the dorsal tubercles on metatarsals II and III Scale bars = 20 mm

Fig. 15 Cladogram of Mesozoic birds showing interrelationships within the Confuciusornithiformes The strict consensus tree is built from 144 most parsimonious trees (Length = 1200; Consistency index = 0.324; Retention index = 0.673) that were found in the present phylogenetic analysis Bootstrap and Bremer values are indicated in bold and italic fonts near the corresponding nodes, respectively

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