柴达木盆地上新世伍氏献文鱼(Cyprinidae: Schizothoracinae)形态学再研究

doi:10.19615/j.cnki.2096-9899.211026

摘要/Abstract

摘要：

柴达木盆地是研究青藏高原古环境和动物演化历史的一个关键区域。为了适应上新世时期柴达木盆地环境的干旱化,化石裂腹鱼类伍氏献文鱼(Hsianwenia wui)演化出了异常粗大的骨骼。然而,人们对于这种骨骼增粗现象(pachyostosis)的性质仍知之甚少。为了进一步认识这一特征及其生理学机制,对伍氏献文鱼进行了详尽的形态解剖学工作,并识别出了一些新的信息：脑腔前部分叉的嗅束通道、韦伯氏器上发达的第3髓上骨、第5尾前椎支持众多尾鳍短鳍条以及第2尾前椎上一个附加的髓弓(棘)。此外,发现献文鱼骨骼的增粗存在差异性：该现象仅见于内骨骼,外骨骼(膜质骨)一般未见增粗;在成组增粗的内骨骼(如肌间骨：上髓弓小骨和上肋小骨)中,腹侧骨骼较背侧增粗更为明显。伍氏献文鱼匙骨后缘有一个显著而独特的位于胸鳍上方的“肩突”(humeral process), 对比现生鲤科鱼类咀嚼活动中相关肌肉和骨骼的联动关系,认为这个“肩突”的出现与伍氏献文鱼咽颌骨骼(第五角鳃骨)增粗以及研磨坚硬的食物有关。

关键词: 柴达木盆地, 上新世, 伍氏献文鱼, 形态学, 骨肥厚, 咀嚼系统和匙骨“肩突”

Abstract:

The Qaidam Basin is a key area for understanding the paleoenvironmental and faunal history of the Tibetan Plateau. The fossil schizothoracine fish, Hsianwenia wui, evolved extraordinarily thickened bones to adapt to the aridification of the Qaidam Basin during the Pliocene. However, the nature of the bone thickening itself remains elusive. To promote the further investigation of the physiological mechanism of the pachyostosis and the phylogenetic interrelationships of Hsianwenia and all relevant cyprinids, here we present a comprehensive morphological study of Hsianwenia. We have new information on the anterior part of the cranial cavity, a large supraneural 3 in the Weberian apparatus, numerous procurrent caudal fin rays supported by the preural centrum (Pu) 5, and a neural arch on Pu2. We also find the differentiated pattern of the bone-thickening: the pachyostosis exists in the endoskeleton but not in the dermal skeleton; it is more obvious in ventral bones than in dorsal ones, when the thickening is present in the dorsally and ventrally grouped endoskeletal bones (e.g., the epineural and epipleural intermuscular bones). Considering the integrity of musculoskeletal system manipulating the chewing activities, we suspect that the thickened pharyngeal jaws and the hard food processing might be associated with the unique hind protrusion (cleithral “humeral” process) of the dermal pectoral girdle of Hsianwenia.

Key words: Qaidam Basin, Pliocene, Hsianwenia wui, morphology, differential skeletal thickening, chewing system and cleithral humeral process

中图分类号:

Q915.862

毕黛冉, 吴飞翔, 王宁, 张弥曼, 房庚雨. 柴达木盆地上新世伍氏献文鱼(Cyprinidae: Schizothoracinae)形态学再研究. 古脊椎动物学报, 2022, 60(1): 1-28.

BI Dai-Ran, WU Fei-Xiang, WANG Ning, CHANG Mee-Mann, FANG Geng-Yu. Revisit of Hsianwenia wui (Cyprinidae: Schizothoracinae) from the Pliocene of Qaidam Basin. Vertebrata Palasiatica, 2022, 60(1): 1-28.

图/表 13

Fig. 1 Hsianwenia wui Chang et al., 2008 from the Pliocene of Qaidam Basin A. photograph of holotype (IVPP V 15244); B. skeletal restoration. Anterior facing right Arrows in the caudal fin in B indicate the outermost upper and lower principal rays. Scale bar equals 20 mm

Table 1 Measurements and meristics (ratio) of Hsianwenia wui (IVPP V 15244) (mm)

Total length	460-510
Standard length	417.7
Body depth	77.4
Head length	109.2
Head depth	63.0
Snout length	17.8
Postorbital length	63.9
Distance between snout tip and dorsal fin origin	231.3
Distance between dorsal fin and caudal fin base	186.4
Distance between anal fin origin and pelvic fin insertion	99.8
Distance between anal fin origin and caudal fin base	75.1
Distance between pectoral fin insertion and pelvic fin insertion	103.5
Dorsal fin base length	46.4
Anal fin base length	29.1
Caudal peduncle length	47.0
Caudal peduncle depth	24.3
Pectoral fin length	84.5
Pelvic fin length	61.8
Standard length/body depth	5.4
Head length/head depth	1.7
Standard length/head length	3.8
Caudal peduncle length/caudal peduncle depth	1.9

Fig. 2 Hsianwenia wui Chang et al., 2008 from the Pliocene of Qaidam Basin, skull, pectoral girdle and anterior part of pectoral fins (holotype IVPP V 15244) A. photograph; B. line drawing Abbreviations: aCh. anterior ceratohyal; An. anguloarticular; Apa. autopalatine; br. branchiostegal rays; Cl. cleithrum; De. dentary; Ecpt. ectopterygoid; Enpt. entopterygoid; Epo. epioccipital; fna. foramen of hyoid artery and facial nerve; Fr. frontal; Hh. hypohyal; Hy. hyomandibular; Io2-3. infraorbital 2-3; Iop. interopercle; Le. lateral ethmoid; lp2. lateral process of the second vertebral centrum; Me. mesethmoid; Mx. maxilla; Mpt. metapterygoid; ocpip. origin of m. pharyngo-cleithralis internus posterior; oos. outer arm of the os suspensorium; Op. opercle; Pa. parietal; pf. pectoral fin; Pmx. premaxilla; Pop. preopercle; Pt. posttemporal; Pto. pterotic; Qu. quadrate; rb. rib; Scl. supracleithrum; Soc. supraoccipital; Sop. subopercle; Spo. sphenotic; T. tripus. Scale bar equals 10 mm. Anterior facing right

Fig. 3 Hsianwenia wui Chang et al., 2008 from the Pliocene of Qaidam Basin, skull of specimen IVPP V 15012 A. photograph; B. line drawing Abbreviations: Cb. ceratobranchial; gr. gill rakers; l. left; pb. pharyngeal bone; r. right For other abbreviations see Fig. 2. Scale bar equals 10 mm. Anterior facing right

Fig. 4 Hsianwenia wui Chang et al., 2008 from the Pliocene of Qaidam Basin, kinethmoid (IVPP V15306-1) (A-C) and orbitosphenoid (V 15306-2) (D-G) A. front view; B. back view; C. lateral view of kinethmoid. D. dorsal view; E. ventral view; F. front view; G. back view of orbitosphenoid. Dorsal facing up (A-C, F-G), figures marked with 1 are photographs, and 2 are line drawings Abbreviations: d. depression for ligament connecting dorsal processes of premaxillae; fLe. facet originally abutting lateral ethmoid; fPts. facet originally abutting pterosphenoid; fva. vascular foramina; gI. Grooves probably for passages of the olfactory tracts; p. process to connective tissue linking mesethmoid and vomer; rtel. recess for the telencephalon. The scale bars equal 1 mm in A-C, and equal 2 mm in D-G

Fig. 5 Hsianwenia wui Chang et al., 2008 from the Pliocene of Qaidam Basin, disarticulated dentary and anguloarticular A, B. photo (A1, B1) and drawing (A2, B2) of a right dentary (IVPP V 15306.1) in median (A) and lateral (B) views; C, D. photo (C1, D1) and drawing (C2, D2) of a left anguloarticular (V 15306.2) in lateral (C) and median (D) views Abbreviations: cp. coronoid process; fVmd. foramen for mandibular branch of trigeminal nerve; fVIImd. foramen for mandibular branch of facial nerve; gac. groove accommodating anguloarticular and Meckel’s cartilage; j. joint facet for counterpart dentary (mandibular symphysis); jRa. joint facet for retroarticular; Mco. the origin of the Meckel’s cartilage; msc. mandibular sensory canal; sQu. socket for the articular head of quadrate. Dorsal facing up. All scale bars equal 2 mm

Fig. 6 Hsianwenia wui Chang et al., 2008 from the Pliocene of Qaidam Basin, disarticulated opercular series bones A. left preopercle, medial view, in IVPP V 15245.1; B. the third branchiostegal ray in V 15245.2; C. the second or first branchiostegal in V 15245.2. Figures marked with 1 are photographs, and 2 are line drawings. Anterior facing right in A, and left in B-C, dorsal facing up Scale bars equal 20 mm in A, and equal 5 mm in B and C

Fig. 7 Hsianwenia wui Chang et al., 2008 from the Pliocene of Qaidam Basin, left hyomandibular (IVPP V 15306.3), pharyngeal bone and teeth in medial view (V 15244) and urohyal (V 28581) A, B. hyomandibular in medial (A) and lateral (B) views. Dorsal facing up. C. pharyngeal bone in medial view; D, E. urohyal in dorsal (D) and posterior (E) views, anterior facing left in D Abbreviations: ae. anterior edentulous process; fna. foramen of hyoid artery and facial nerve; hp. horizontal ventral plate; jn. dorsal joint processes to neurocranium; jOp. joint process to opercle; vp. longitudinal vertical plate Scale bars equal 10 mm Figures marked with 1 are photographs, and 2 are line drawings

Fig. 8 Hsianwenia wui Chang et al., 2008 from the Pliocene of Qaidam Basin, vertebrae A. abdominal vertebrae (IVPP V 15245.1); B. abdominal vertebrae (V 15244); C. caudal vertebrae (V 15245.1); D. caudal vertebrae (V 15012); E, F. left os suspensorium (V 15306) in front (E) and back (F) views Anterior facing left in A, C and F; anterior facing right in B, D, E. Dorsal facing up in all figures Figures marked with 1 are photographs and those marked with 2 are line drawings Abbreviations: do. distal end of outer arm of os suspensorium; En. epineural; Epl. epipleural; hs. haemal spine; ios. inner arm of os suspensorium (broken); lp2. lateral process of the second vertebral centrum; ns. neural spine; oos. the outer arm of the os suspensorium; pv. parapophysis; rb. rib; Sn3. supraneural 3; uh. upper head of os suspensorium; vc4, vc38. the 4th and 38th centra. Scale bars equal 10 mm

Fig. 9 Hsianwenia wui Chang et al., 2008 from the Pliocene of Qaidam Basin, intermuscular bones A. an epineural, IVPP V 15306.20; B. an epipleural, V 15012. Scale bars equal 5 mm Figures marked with 1 are photographs, and 2 are line drawings

Fig. 10 Hsianwenia wui Chang et al., 2008 from the Pliocene of Qaidam Basin, caudal skeleton and caudal fin (holotype IVPP V 15244) A. photograph; B. line drawing Abbreviations: cc. compound centrum; Ep. epural; Epl. epipleural; hs. haemal spine; H1-H5. hypural 1-5; ns. neural spine; Ph. parhypural; Pls. pleurostyle; Pu2, 5. preural centra 2, 5; Ur. Uroneural Arrow pointing to the outermost principal rays in upper and lower lobes, scale bar equals 5 mm

Fig. 11 Hsianwenia wui Chang et al., 2008 from the Pliocene of Qaidam Basin, median fins, paired fins and girdles A. dorsal fin (holotype, IVPP V 15244); B. anal fin (V 15244); C. anal fin (V 15012); D. pectoral fin (V 15244); E. pelvic fin V 15244); F. left basipterygium (V 15245.1). Anterior facing right, dorsal facing up Abbreviations: afs. anal fin stay; ap1. first anal fin pterygiophore; Bp. basipterygium; dp1. first dorsal fin pterygiorphore; fpr. facet for articulation of pelvic radials; ip. ischiac process; jp. joint process; Sn. supraneural. Scale bars equal 5 mm in F and 10 mm in the rest

Fig. 12 Musculoskeletal correlation of pharyngeal jaws and pectoral girdle (cleithrum) A. general cyprinid-type chewing system, Cyprinus carpio (redrawn from Sibbing, 1982); B. tentative restoration of chewing system of Hsianwenia wui based on A. Anterior facing left Abbreviations: ac. anterior cartilage; Cbr4. ceratobranchial 4; Cl. cleithrum; m.cpe. m. pharyngo-cleithralis externus; m.cpia. m. pharyngo-cleithralis internus anterior; m.cpip. m. pharyngo-cleithralis internus posterior; m.hpa. hypaxial trunk muscle; pb. pharyngeal bone; pcp. posterior cleithral process; pf. pectoral fin; Scl. supracleithrum. Drawings not to scale

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