燕辽生物群的两件蛙嘴翼龙类新标本及对蛙嘴翼龙类头骨的新解释

doi:10.19615/j.cnki.2096-9899.260411

摘要/Abstract

摘要：

蛙嘴翼龙是一类从中侏罗世到早白垩世时期分布于欧亚大陆的小型非翼手龙类翼龙。其主要的鉴别特征是短而宽的头骨，这一形态与所有其他翼龙类群都不同。由于保存的原因，蛙嘴翼龙类的头骨结构长期以来一直都存在争议。详细描述了两件来自燕辽生物群髫髻山组的蛙嘴翼龙类标本，并对其中一件标本中保存相对完好的头骨进行了计算机断层扫描与重建。结果显示，该头骨存在一个与大部分翼龙都不相同的独特的骨骼结构，其功能可能类似其他爬行类的眶上骨。基于对该标本头骨的研究，对之前未详细描述的其他蛙嘴翼龙类头骨进行了修订，最终重建了一个新的蛙嘴翼龙类头骨复原图。同时还对蛙嘴翼龙类的头后骨骼部分进行了更为全面的比较分析和讨论。

关键词: 燕辽生物群, 髫髻山组, 翼龙, 蛙嘴翼龙类, 古尾翼龙属

Abstract:

Anurognathids are a clade of small, early-diverging non-pterodactyloid pterosaurs distributed across Eurasia from the Middle Jurassic to the Early Cretaceous. Their defining characteristic is a short and broad skull, a morphology distinct from all other pterosaur clades. Due to limited preservation, the cranial anatomy of this group has long been difficult to understand in detail. In this study, we provide a detailed description of two anurognathid specimens from the Tiaojishan Formation of the Yanliao Biota. The relatively well-preserved skull of one specimen was reconstructed through computed tomographic scanning. This reveals a unique skull element, possibly functionally analogous to the supraorbital bone of varanid lizards. The CT reconstruction offers new insights into the anurognathid skull. Based on these findings, we revised the skulls of previously described taxa, ultimately integrating them into a new reconstruction for this clade. We present a detailed comparative analysis and discussion of the postcranial osteology within Anurognathidae.

Key words: Yanliao Biota, Tiaojishan Formation, Pterosaur, Anurognathid, Cascocauda

中图分类号:

Q915.864

佟仕达, 蒋顺兴, 程心, 汪筱林. 燕辽生物群的两件蛙嘴翼龙类新标本及对蛙嘴翼龙类头骨的新解释. 古脊椎动物学报, 2026, 64(2): 191-220.

TONG Shi-Da, JIANG Shun-Xing, CHENG Xin, WANG Xiao-Lin. Two anurognathid pterosaur specimens from the Yanliao Biota and a new interpretation of anurognathid skulls. Vertebrata Palasiatica, 2026, 64(2): 191-220.

图/表 11

Fig. 1 Overview photograph (A) and line drawing (B) of the specimen IVPP V23669 from the Middle-Upper Jurassic Qinglong, Hebei Abbreviations: cd. caudal vertebra; co. coracoid; cr. cervical rib; cv. cervical vertebra; dr. dorsal rib; f. femur; fr. frontal; ga. gastralia; h. humerus; il. ilium; is. ischium; j. jugal; l. left; m. mandible; mcIV. metacarpal IV; mtI-V. metatarsal I-V; mx. maxilla; pa. parietal; pm. premaxilla; po. postorbital; ppu. prepubis; PI-IV. pedal digits I-IV; PV1 and 2. phalanges 1 and 2 of pedal digit V; r. right; rd. radius; sc. scapula; ti. tibia; u. ulna; wp1-2. wing phalanges 1-2. Scale bars equal 10 mm

Fig. 2 The cranium of specimen IVPP V23669 from the Middle-Upper Jurassic Qinglong, Hebei A-C. the digital reconstruction (A), line drawing (B), and photograph (C) of the skull in ventral view; D, E. the digital reconstruction (D) and line drawing (E) in dorsal view Abbreviations: ecp. ectopterygoid; fr. frontal; hy. hyoid; j. jugal; m. mandible; mx. maxilla; n. nasal; pa. parietal; pal. palatine; pm. premaxilla; po. postorbital; prf. prefrontal; ps. parasphenoid; pt. pterygoid; q. quadrate; scr. sclerotic ring; sq. squamosal; t. tooth; v. vomer. Scale bars equal 5 mm

Fig. 3 Digital reconstruction of cranial elements of specimen IVPP V23669 in dorsal view (only J in ventral view) from the Middle-Upper Jurassic Qinglong, Hebei A. premaxillae; B. left and right maxillae; C. left and right prefrontals; D. right postorbital; E. right jugal; F. right quadrate; G. left and right palatines; H. left and right vomers/pterygoids; I. left and right hyoids; J. parasphenoid; K. left and right ectopterygoids Abbreviations: apm. ascending process of the maxilla; apr. apex of prefrontal; fpp. frontal process of prefrontal; jpe. jugal process of ectopterygoid; mpp. median process of premaxilla; ppe. pterygoid process of ectopterygoid; ppj. postorbital process of jugal; ppp. posterior process of prefrontal; sqp. squamosal process of postorbital. Scale bars equal 2 mm (A, C, D, E, F, H and J) and 5 mm (B, G, I and K)

Fig. 4 The digital reconstruction of the teeth in specimen IVPP V23669 in dorsal view from the Middle-Upper Jurassic Qinglong, Hebei Abbreviations: dt. dentary tooth; l. left; mxt. maxillary tooth; pmt. premaxillary tooth; r. right Scale bar equals 5 mm

Fig. 5 Photograph (A) and line drawing (B) of the close-up of pelvic girdles of IVPP V23669 from the Middle-Upper Jurassic Qinglong, Hebei Abbreviations: cd. caudal vertebrae; ch. chevron; f. femur; ga. gastralia; il. ilium; is. ischium; l. left; ppu. prepubis; pu. pubis; r. right; sr. sacral rib. Scale bars equal 5 mm

Fig. 6 Photograph (A) and line drawing (B) of the specimen IVPP V26040 from the Middle-Upper Jurassic Linglongta, Liaoning Abbreviations: cd. caudal vertebrae; co. coracoid; cr. cervical rib; cv. cervical vertebrae; dr. dorsal rib; f. femur; fi. fibula; ga. gastralia; h. humerus; il. ilium; is. ischium; l. left; mcIV. metacarpal IV; mtI-V. metatarsal I-V; p. pteroid; pu. pubis; PI-IV. pedal digits I-IV; PV1 and 2. phalanges 1 and 2 of pedal digit V; q. quadrate; r. right; rd. radius; sc. scapula; ti. tibia; u. ulna; wp1-4. wing phalanges 1-4 Scale bars equal 10 mm

Fig. 7 Close-up photographs (A, C, E) and line drawings (B, D, F) of caudal vertebrae, sternum and pelvic girdle of IVPP V26040 from the Middle-Upper Jurassic Linglongta, Liaoning A, B. completely preserved tail with zygapophyses and chevrons; C, D. sternum exposed in ventral view; E, F. pelvic girdle. Abbreviations: ch. chevron; co. coracoid; f. femur; il. ilium; l. left; is. ischium; pu. pubis; r. right; st. sternum; z. zygapophyses. Scale bars equal 5 mm (A, B, E and F) and 1 mm (C, D)

Fig. 8 The line drawings of the skulls of other anurognathids A. Dendrorhynchoides (GMV 2128); B. Luopterus (JZMP-04-07-3); C. Jeholopterus (IVPP V12705) (the light grey denotes areas preserved as impressions on V12705B, dark grey denotes areas represented by preserved bone, and dashed lines indicate regions where the bone shape is discernible beneath overlying elements) Abbreviations: ang. angular; bp. basipterygoid; cv. cervical vertebrae; d. dentary; ecp. ectopterygoid; fr. frontal; hy. hyoid; j. jugal; mx. maxilla; pa. parietal; pal. palatine; pm. premaxilla; po. postorbital; prf. prefrontal; ps. parasphenoid; pt. pterygoid; q. quadrate; qj. quadratojugal; sa. surangular; scr. sclerotic ring; sq. squamosal; t. tooth; v. vomer. Scale bars equal 2 mm (B) and 5 mm (A, C)

Fig. 9 Reconstruction of anurognathids, redrawn and modified from Bennett (2007) and Dalla Vecchia (2022) Blue areas indicate newly added elements; yellow areas indicate revised regions A. dorsal view; B. ventral view; C. right lateral view. Abbreviations: ecp. ectopterygoid; fr. frontal; j. jugal; m. mandible; mx. maxilla; n. nasal; pa. parietal; pal. palatine; pm. premaxilla; po. postorbital; prf. prefrontal; ps. parasphenoid; pt. pterygoid; q. quadrate; qj. quadratojugal; scr. sclerotic ring; sq. squamosal; v. vomer Not drawn to scale

Table 1 Tooth counts and the measurements of teeth and skulls for anurognathids (mm)

Taxon	Specimen	pmt	mxt	dt	pmt length	skull length
Anurognathus ammoni	BSP 1922 I 42	3	6	>3	2.7	~27
Anurognathus ammoni	SMNS 81928	-	-	-	-	15.3
Batrachognathus volans	PIN 52-2	3	8	-	3.2	48
Dendrorhynchoides curvidentatus	GMV 2128	>2	>2	>3	3.03	>17.85
Jeholopterus ningchengensis	IVPP V12705	3	>3	>1	3.85	28
Jeholopterus ningchengensis	CAGS-Z070	3	>3	-	-	21.1
Luopterus mutoudengensis	JZMP-04-07-3	-	-	>4	-	24.75
Vesperopterylus lamadongensis	BMNHC-PH-001311	-	-	-	3.51	31.8
Sinomacrops bondei	JPM-2012-001	-	9	-	-	>16.26
Cascocauda rong	NJU-57003	3	>3	-	>2.87	22.2
Cascocauda rong	IVPP V23669	3	4	7	3.27	>19
indeterminate	IVPP V16728	-	-	-	3.1	-

Fig. 10 The line drawings of anurognathid humeri, the arrows indicate the deltopectoral crests and the opposing structure the ulnar crests A. left humerus of Anurognathus ammoni (SMNS 81928); B. right humerus of A. ammoni (SMNS 81928); C. left humerus of Sinomacrops bondei (JPM-2012-001); D. left humerus of IVPP V23669; E. right humerus of IVPP V23669; F. right humerus of Luopterus mutoudengensis (JZMP-04-07-3); G. left humerus of Cascocauda rong (NJU-57003); H. right humerus of C. rong (NJU-57003); I. left humerus of IVPP V26040; J. left humerus of Dendrorhynchoides curvidentatus (GMV 2128); K. left humerus of A. ammoni (BSP 1922 I 42); L. left humerus of Batrachognathus volans (PIN 52-2); M. left humerus of Jeholopterus ningchengensis (CKGP 19960828) (based on Gao et al., 2009); N. left humerus of J. ningchengensis (IVPP V12705); O. right humerus of the holotype specimen of Vesperopterylus lamadongensis (BMNHC-PH-001311) Scale bars equal 5 mm

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