赫氏近鸟龙(兽脚类恐龙：副鸟类)头骨形态对鸟类头骨演化的新启示

doi:10.19615/j.cnki.2096-9899.241225

摘要/Abstract

摘要：

无论从何种维度，恐龙—鸟类的演化都是地球生命历史中最吸引人的一次演化事件，其中涉及大量形态学和生态学特征的改变。相较于头后骨骼形态，现有的研究对鸟类头骨的早期演化所取得的认识相对有限，这主要受限于保存较好的早期鸟翼类头骨化石材料的稀少。近鸟龙是目前已知最早的副鸟类(距今约1.6亿年)——副鸟类是指包含所有现代鸟类，但不包括尾羽龙类或者耀龙类的最广义类群。已知的近鸟龙类化石多达百余件，使得该类群成为研究非鸟兽脚类恐龙—鸟类演化过程中形态变化的最理想对象。然而目前有关近鸟龙类的头骨形态特征存在大量未知的内容。基于一件保存较好的近鸟龙类化石，详细描述了该类群头骨的形态结构，包括此前知之甚少的区域(例如颞区和颚区)。研究显示出近鸟龙保留了非鸟恐龙所具有的原始的双弓型非可动性的头骨形态。同时，近鸟龙头骨还呈现出与驰龙类、伤齿龙类，以及原始鸟翼类分别相似的局部形态特征，揭示了模块化演化深刻影响鸟翼类头骨的早期演化。

关键词: 近鸟龙, 鸟翼类, 头骨, 可动性, 颚骨, 鳞状骨

Abstract:

The origin of birds from theropod dinosaurs, by any measures, is the most eye-catching evolutionary transition in the history of life, which encompasses numerous extensive morphological and biological changes. Compared to postcranium, little progress has been made regarding the evolutionary assemblage of the birds’ skull, because of few detailed early records of cranial materials of stem lineages. Anchiornis is the oldest known record of the Paraves (~160 Ma), the most inclusive clade that contains all living birds but not Caudipteryx or Epidexipteryx. With hundreds of known specimens, Anchiornis constitutes an ideal taxon for investigating morphological modifications across the theropod-bird transition, but its cranial morphology remains enigmatic. Here we present in-depth description of the cranial morphology of Anchiornis based on three-dimensional reconstruction of a well-preserved specimen, including elements from the temporal and palatal regions that are poorly recognized previously. Our study shows that Anchiornis retains the plesiomorphic dinosaurian condition in having a diapsid akinetic skull. The mixture of cranial characters, shared with dromaeosaurids, troodontids, and stemward avialans, present in Anchiornis demonstrates the complex history of early avialan cranial evolution.

Key words: Anchiornis, Avialae, cranium, kinesis, palatine, squamosal

中图分类号:

Q915.865

王敏, 王孝理, 郑晓廷, 周忠和. 赫氏近鸟龙(兽脚类恐龙：副鸟类)头骨形态对鸟类头骨演化的新启示. 古脊椎动物学报, 2025, 63(1): 20-42.

WANG Min, WANG Xiao-Li, ZHENG Xiao-Ting, ZHOU Zhong-He. Cranial anatomy of Anchiornis huxleyi (Theropoda: Paraves) sheds new light on bird skull evolution. Vertebrata Palasiatica, 2025, 63(1): 20-42.

图/表 9

Fig. 1 Photograph of Anchiornis huxleyi (STM 0-47) from Jianchang, Liaoning Scale bar = 10 cm

Fig. 2 Photograph of the skull of Anchiornis huxleyi (STM 0-47) from Jianchang, Liaoning Scale bar = 10 mm

Fig. 3 Digital reconstitution of the skull of Anchiornis huxleyi (STM 0-47) from Jianchang, Liaoning in right (A) and left (B) lateral views showing cranial anatomy Abbreviations: hy. hyoid; lan. left angular; lde. left dentary; lec. left ectopterygoid; lfr. left frontal; lju. left jugal; llc. left lacrimal; lma. left maxilla; lna. left nasal; lpa. left palatine; lpm. left premaxilla; lpo. left postorbital; lqj. left quadratojugal; lqu. left quadrate; ls. left squamosal; lsp. left splenial; lsu. left surangular; pa. prearticular; par. parietal; pr. parasphenoid rostrum; ran. right angular; rde. right dentary; rec. right ectopterygoid; rfr. right frontal; rlc. right lacrimal; rma. right maxilla; rna. right nasal; rpa. right palatine; rpm. right premaxilla; rpo. right postorbital; rqj. right quadratojugal; rqu. right quadrate; rsu. right surangular; vo. vomer. Scale bars = 10 mm

Fig. 4 Anatomy of facial elements of Anchiornis huxleyi (STM 0-47) from Jianchang, Liaoning Digital reconstruction of the left premaxilla (A), right maxilla (B), left lacrimal (C, D), and left nasal (E, F) A-C, E. lateral views; D, F. medial views. The blue arrowhead indicates the recess at the juncture of the anterior and ventral processes, and the orange arrowhead denotes the lateral fossa on the ventral process Abbreviations: afl. articular facet for lacrimal; apl. anterior process of lacrimal; ib. interfenestral bar; jm. jugal process of maxilla; mf. maxillary fenestra; mp. maxillary process; np. nasal process; pf. promaxillary fenestra; pm. premaxillary process; pn. premaxillary process of nasal; pp. promaxillary pila; ppl. posterior process of lacrimal; vn. ventral process of nasal; vpl. ventral process of lacrimal. Scale bars = 5 mm

Fig. 5 Anatomy of the temporal elements of Anchiornis huxleyi (STM 0-47) from Jianchang, Liaoning Digital reconstruction of the left postorbital (A), left squamosal (B, C), and right jugal and quadratojugal (D, E) A, B, E. lateral views; C, D. medial views Abbreviations: afp. articular facet for postorbital; afq. articular facet for quadrate; fpp. frontal process of postorbital; jpp. jugal process of postorbital; jpq. jugal process of quadratojugal; mpj. maxillary process of jugal; pas. parietal process of squamosal; ppj. postorbital process of jugal; pps. postorbital process of squamosal; qps. quadratojugal process of squamosal; spp. squamosal process of postorbital; spq. squamosal process of quadratojugal. Scale bars = 5 mm

Fig. 6 Anatomy of the quadrate of Anchiornis huxleyi (STM 0-47) from Jianchang, Liaoning A-D. right quadrate in lateral (A), medial (B), posterior (C), and ventral (D) views; E, F. left quadrate in lateral (E) and medial (F) views Abbreviations: afq. articular facet for quadratojugal; lc. lateral condyle; mc. medial condyle; mp. mandibular process; orp. orbital process; otp. otic process; qr. quadrate ridge. Scale bars = 5 mm

Fig. 7 Anatomy of the palatal elements of Anchiornis huxleyi (STM 0-47) from Jianchang, Liaoning A, B. right pterygoid in medial (A) and lateral (B) views; C, D. left palatine in dorsal (C) and ventral (D) views; E, F. right ectopterygoid in dorsal (E) and ventral (F) views; G. parasphenoid rostrum in ventrolateral view; H. possible right vomer in dorsal view The blue arrowheads in (C) denote the dorsal depressions, and the green arrowhead in (G) denotes the concave ventral surface of the parasphenoid rostrum Abbreviations: afp. articular facet for basipterygoid process; chp. choanal process; jue. jugal process of ectopterygoid; jup. jugal process; mxp. maxillary process; pfe. pterygoid flange of ectopterygoid; plr. palatine ramus; pte. pterygoid process of ectopterygoid; ptp. pterygoid process of palatine; qur. quadrate ramus. Scale bars = 5 mm

Fig. 8 Anatomy of the lower jaw elements of Anchiornis huxleyi (STM 0-47) from Jianchang, Liaoning A, B. left dentary in medial (A) and lateral (B) views; C. left splenial in medial view; D, E. left post-dentary elements in medial (D) and lateral (E) views; F. right post-dentary elements in lateral view Abbreviations: ang. angular; emf. external mandibular foramen; saf. surangular foramen; san. Surangular Scale bars = 5 mm

Fig. 9 Evolutionary tree showing the position of Anchiornis

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