广西下泥盆统的肉鳍鱼类化石

doi:10.19615/j.cnki.2096-9899.241226

摘要/Abstract

摘要：

记述了广西南宁下泥盆统莲花山组和那高岭组中发现的早期肉鳍鱼类化石，材料包括一件斑鳞鱼(Psarolepis)的左侧颊板骨，一件杨氏鱼(Youngolepis)的后顶甲，一件奇异鱼(Diabolepis)的颅顶甲和一枚蝶柱鱼(Styloichthys)鳞片。其中颊板骨标本代表了目前已知最晚的斑鳞鱼化石记录，奇异鱼和蝶柱鱼标本则是这两个属在云南曲靖以外的首次报道。这些材料丰富了徐家冲组合的生物面貌，为广西、云南和越南北部下泥盆统的地层对比提供了重要化石证据。根据最新的牙形石证据，广西南宁地区的徐家冲组合可能最早在晚洛霍考夫期就已出现，比其他地区该组合的出现时间更早。广西南宁的莲花山组与那高岭组的过渡地层可能与曲靖地区的桂家屯组时代相当。同时，南宁地区的鱼类化石标本较云南曲靖地区的有明显大型化趋势，可能与当时南宁地区鱼类的生活环境中氧含量高有关。

关键词: 广西南宁, 下泥盆统, 洛霍考夫期, 徐家冲组合, 莲花山组, 那高岭组, 肉鳍鱼类

Abstract:

Here we report a left cheek plate of Psarolepis, a postparietal shield of Youngolepis, a skull of Diabolepis, and a scale of Styloichthys from the Lianhuashan and Nahkaoling formations (Lochkovian, Lower Devonian) of Nanning, Guangxi. This marks the first report of Diabolepis and Styloichthys beside Qujing, Yunnan, and the latest occurrence of Psarolepis to date. The fossil community displays significant similarities to the Xujiachong Assemblage, and provides new data for the Lower Devonian stratigraphic correlation between southwestern China and northern Vietnam. Given the latest dating constraint based on the conodont evidence, we regard that the Xujiachong Assemblage has a much longer range than previously supposed, extending from the latest Lochkovian to the end of Pragian. We propose that the transition of the Nahkaoling and Lianhuashan formations in Nanning might correspond to the Guijiatun Formation in Qujing. The relatively large size of fish individuals from Guangxi is probably attributed to the increase in the oxygen content of the ocean.

Key words: Nanning,Guangxi, Lower Devonian, Lochkovian, Xujiachong Assemblage, Lianhuashan Formation, Nahkaoling Formation, Sarcopterygians

中图分类号:

Q915.862

李懋坤, 崔心东, 朱敏, 乔妥. 广西下泥盆统的肉鳍鱼类化石. 古脊椎动物学报, 2025, 63(1): 1-19.

LI Mao-Kun, CUI Xin-Dong, ZHU Min, QIAO Tuo. Sarcopterygians from the Lochkovian (Lower Devonian) of Nanning, Guangxi, China. Vertebrata Palasiatica, 2025, 63(1): 1-19.

图/表 7

Fig. 1 Locality and geological setting of the fossil sites A. location and geologic map of the fossil locality (based on the open geological data in the National Geological Archives of China: http://www.ngac.org.cn/Map); B. stratigraphic sequence of the Lower Devonian in the Nanning-Liujing district (modified after Kuang et al., 1989; Lu et al., 2023) ① Xianhu (22°48'50"N; 108°28'55"E); ② Wuxiangling (22°45'49"N; 108°17'52"E); Є. Cambrian;D1l. Lianhuashan Formation; D1n. Nahkaoling Formation; D1y. Yukiang Formation;D1m-D2. Lower Devonian Moding Formation and the Middle Devonian; D3. Upper Devonian;C. Carboniferous; K. Cretaceous; E. Paleogene; Q. Quaternary

Fig. 2 Photograph (A) and interpretative drawing (B) of Psarolepis sp. (IVPP V20340) Abbreviations: fo1-fo3. foramina; grp. groups of pores; Mx. maxillary; pc. preopercular canal; Pop. preopercular

Fig. 3 Comparison of osteichthyans cheek plates A. Mimipiscis toombsi (modified from Choo, 2012); B. Kenichthys campbelli (modified from Zhu and Ahlberg, 2004); C. Youngolepis sp. (modified from Chang, 1991; Cui et al., 2022);D. Guiyu oneiros (modified from Zhu et al., 2009); E. Psarolepis romeri (modified from Zhu et al., 1999);F. Psarolepis sp. (IVPP V20340) Abbreviations: fo1-fo3. foramina; grp. groups of pores; Mx. maxillary; pc. preopercular canal;pl. pit-line; Pop. preopercular

Fig. 4 Photograph (A), interpretative drawing (B), and 3D virtual model (C) of the scale of Styloichthys sp. (IVPP V33256) Abbreviations: e.a. anterior edge; e.d. dorsal edge; e.p. posterior edge; f.d. depressed field; p. peg; po. pores; r. ridges

Fig. 5 Photograph and comparison of postparietal shields in dipnomorphs A-B. Youngolepis sp. (IVPP V33255): A. photograph, B. reconstruction; C-D. Y. praecursor: C. V6553,D. V6183 (modified from Chang, 1982); E. Powichthys thorsteinssoni (modified from Jessen, 1975);F. Arquatichthys porosus (modified from Lu and Zhu, 2019) Abbreviations: grp. groups of pores; mpl, middle pit-line; ppl. posterior pit-line; Pp. postparietal;St. supratemporal; Ta. tabular

Fig. 6 Photograph and reconstructions of skull roofs and endocast of Diabolepis speratus A, B, D, E. IVPP V33254: A. photograph, B. interpretive drawing, D. 3D virtual model of recognizable structures of the endocast, E. interpretive drawing in lateral view;C. interpretive drawing of IVPP V7237 (modified from Chang and Yu, 1984) Abbreviations: c.pi. pineal canal; mpl. middle pit-line; nc. nasal capsule; om. orbital margin; pi. pineal;ppl. posterior pit-line; soc. supraorbital canal; soc.p. pores for supraorbital canal

Fig. 7 Dating and correlation of the Early Devonian stratigraphy from four selected section areas in South China Block Data are from Liao et al., 1978, 1979; Ma et al., 2009; Xue et al., 2018; Guo et al., 2019; Qie et al., 2019; Yu et al., 2020; Gai et al., 2022; Lu et al., 2023 Abbreviations: Da. Daliancun Member; Fm. Formation; Hen. Hengxian Member; Liu. Liukankou Member; Mb. Member; Mo. Moding Formation; Pri. Pridoli; Xi. Xishancun Formation; Yu. Yulungssu Formation

参考文献 82

[1]	Ahlberg P E, 1991. A re-examination of sarcopterygian interrelationships, with special reference to the Porolepiformes. Zool J Linn Soc, 103(3): 241-287
[2]	Basden A M, Young G C, Coates M I et al., 2000. The most primitive osteichthyan braincase? Nature, 403: 185-188
[3]	Chang M M, 1982. The Braincase of Youngolepis, a Lower Devonian Crossopterygian from Yunnan, south-western China. Ph. D thesis. Stockholm: Department of Geology, University of Stockholm. 1-113
[4]	Chang M M, 1991. Head exoskeleton and shoulder girdle of Youngolepis. In: Chang M M, Liu Y H, Zhang G R eds. Early Vertebrates and Related Problems of Evolutionary Biology. Beijing: Science Press. 355-378
[5]	Chang M M, 1995. Diabolepis and its bearing on the relationships between porolepiforms and dipnoans. Bull Mus Natl Hist Nat, Sect C, 17(1-4): 235-268
[6]	Chang M M, Yu X B, 1981. A new crossopterygian, Youngolepis praecursor, gen. et sp. nov., from Lower Devonian of eastern Yunnan, China. Sci Sin, 24(1): 89-99
[7]	Chang M M, Yu X B, 1984. Structure and phylogenetic significance of Diabolichthys speratus gen. et sp. nov., a new dipnoan-like form from the Lower Devonian of eastern Yunnan, China. Proc Linn Soc NSW, 107(3): 171-184
[8]	Chang M M, Zhu M, 1993. A new osteolepidid from the Middle Devonian of Qujing, Yunnan. Mem Assoc Aust Palaeontol, 15: 183-198
[9]	Choo B, 2012. Revision of the actinopterygian genus Mimipiscis (=Mimia) from the Upper Devonian Gogo Formation of Western Australia and the interrelationships of the early Actinopterygii. Earth Environ Sci Trans R Soc Edinb, 102(2): 77-104
[10]	Choo B, Zhu M, Qu Q M et al., 2017. A new osteichthyan from the late Silurian of Yunnan, China. PLoS One, 12(3): e0170929
[11]	Cui X D, Friedman M, Qiao T et al., 2022. The rapid evolution of lungfish durophagy. Nat Commun, 13(1): 2390 DOI PMID
[12]	Cui X D, Friedman M, Yu Y et al., 2023. Bony-fish-like scales in a Silurian maxillate placoderm. Nat Commun, 14(1): 7622 DOI PMID
[13]	Cui X D, Qiao T, Peng L J et al., in press. New material of the Early Devonian sarcopterygian Styloichthys changae illuminates the origin of cosmine. J Syst Palaeontol
[14]	Davis S P, Finarelli J A, Coates M I, 2012. Acanthodes and shark-like conditions in the last common ancestor of modern gnathostomes. Nature, 486: 247-250
[15]	Dupret V, Zhu M, Wang J Q, 2009. The morphology of Yujiangolepis liujingensis (Placodermi, Arthrodira) from the Pragian of Guangxi (South China) and its phylogenetic significance. Zool J Linn Soc, 157(1): 70-82
[16]	Forey P L, 1998. History of the Coelacanth Fishes. London: Chapman and Hall. 1-419
[17]	Friedman M, 2007. Styloichthys as the oldest coelacanth: implications for early osteichthyan interrelationships. J Syst Palaeontol, 5(3): 289-343
[18]	Friedman M, Brazeau M D, 2010. A reappraisal of the origin and basal radiation of the Osteichthyes. J Vert Paleont, 30(1): 36-56
[19]	Gai Z K, Yu X B, Zhu M, 2017. The evolution of the zygomatic bone from Agnatha to Tetrapoda. Anat Rec, 300(1): 16-29
[20]	Gai Z K, Lu L W, Zhao W J et al., 2018. New polybranchiaspiform fishes (Agnatha: Galeaspida) from the Middle Palaeozoic of China and their ecomorphological implications. PLoS One, 13(9): e0202217
[21]	Gai Z K, Jiang W Y, Zhao W J et al., 2022. Redescription of the Sanqiaspidae (Galeaspida) from the Lower Devonian of South China and its biostratigraphic significance. Palaeobio Palaeoenv, 2022: 173-191
[22]	Giles S, Darras L, Clement G et al., 2015. An exceptionally preserved Late Devonian actinopterygian provides a new model for primitive cranial anatomy in ray-finned fishes. Proc R Soc B, 282: 20151485
[23]	Guo W, Nie T, Sun Y, 2019. New data on the biostratigraphy of the Early Devonian “Spirifer” tonkinensis brachiopod fauna in South China and adjacent region. Palaeobio Palaeoenv, 99(1): 29-43
[24]	Jarvik E, 1966. Remarks on the structure of the snout in Megalichthys and certain other rhipidistid crossopterygians. Ark Zool, 19: 41-98
[25]	Jarvik E, 1980. Basic Structure and Evolution of Vertebrates, Vol. 1. London: Academic Press. 1-575
[26]	Jessen H L, 1975. A new choanate fish, Powichthys thorsteinssoni n.g., n.sp., from the early Lower Devonian of the Canadian Arctic Archipelago. Colloques Int CNRS, 218: 213-222
[27]	Jessen H L, 1980. Lower Devonian Porolepiformes from the Canadian Arctic with special reference to Powichthys thorsteinssoni Jessen. Palaeontogr Abt A, 167: 180-214
[28]	Kuang G D, Zhao M T, Tao Y B, 1989. The Standard Devonian Section of China:Liujing Section of Guangxi. Wuhan: China University of Geosciences Press. 1-154
[29]	Liao W H, Xu H H, Wang C Y et al., 1978. Division and correlation of Devonian in southwestern China. In: Institute of Geology and Mineral Resources, Chinese Academy of Geological Science ed. Symposium on the Devonian System of South China. Beijing: Geological Publishing House. 193-213
[30]	Liao W H, Xu H K, Wang C Y et al., 1979. On some basic Devonian sections in Southwestern China. In: Nanjing Institute of Geology and Palaeontology ed. Carbonate Biostratigraphy in Southwestern Regions. Beijing: Science Press. 221-249
[31]	Liu S F, 1980. On some arctolepids from the Devonian of Guangxi. Vert PalAsiat, 18(2): 84-88
[32]	Liu S F, 1982. Preliminary note on the Arthrodira from Guangxi, China. Vert PalAsiat, 20(2): 106-114
[33]	Liu S F, 1986. Fossil Eugaleaspid from Guangxi. Vert PalAsiat, 24(1): 1-9
[34]	Liu S F, 1992. On the Lower Devonian antiarchians of Guangxi, China. Vert PalAsiat, 30(3): 210-220
[35]	Liu Y H, 1985. A galeaspid (Agnatha), Antiquisagittaspis cornuta gen. et sp. nov., from Lower Devonian of Guangxi, China. Vert PalAsiat, 23(4): 247-254
[36]	Long J A, 2001. On the relationships of Psarolepis and the onychodontiform fishes. J Vert Paleont, 21(4): 815-820
[37]	Lu J, Zhu M, 2008. An Early Devonian (Pragian) sarcopterygian from Zhaotong, Yunnan, China. Vert PalAsiat, 46(3): 161-170
[38]	Lu J, Zhu M, 2019. The postparietal shield of the Pragian dipnomorph Arquatichthys and its implications for the rhipidistian cranial anatomy. Palaeoworld, 28(4): 543-549
[39]	Lu J, Zhu M, Long J A et al., 2012. The earliest known stem-tetrapod from the Lower Devonian of China. Nat Commun, 3(1): 1160
[40]	Lu J, Zhu M, Ahlberg P E et al., 2016. A Devonian predatory fish provides insights into the early evolution of modern sarcopterygians. Sci Adv, 2(6): e1600154
[41]	Lu J, Giles S, Friedman M et al., 2017. A new stem sarcopterygian illuminates patterns of character evolution in early bony fishes. Nat Commun, 8: 1932 DOI PMID
[42]	Lu J F, Guo W, Yi W et al., 2023. The first discovery of Lochkovian (Lower Devonian) conodonts in central Guangxi, South China and its geological implications. J Paleontol, 97(2): 421-433
[43]	Ma X P, Liao W H, Wang D M, 2009. The Devonian System of China, with a discussion on sea-level change in South China. Spec Publ Geol Soc London, 314(1): 241-262
[44]	Miles R S, 1977. Dipnoan (lungfish) skulls and the relationships of the group: a study based on new species from the Devonian of Australia. Zool J Linn Soc, 61(1-3): 1-328
[45]	P’an K, 1964. Some Devonian and Carboniferous fishes from South China. Acta Palaeontol Sin, 12(1): 139-168
[46]	P’an K, Wang S T, 1978. Devonian Agnatha and Pisces of South China. In: Institute of Geology and Mineral Resources, Chinese Academy of Geological Science ed. Symposium on the Devonian System of South China. Beijing: Geological Publishing House. 298-333
[47]	P’an K, Wang S T, Liu Y P, 1975. The Lower Devonian Agnatha and Pisces from South China. Prof Pap Stratigr Palaeont, 1: 135-169
[48]	P’an K, Wang S T, Gao L D et al., 1978. The Devonian System of South China. In: Institute of Geology and Mineral Resources, Chinese Academy of Geological Science ed. Symposium on the Devonian System of South China. Beijing: Geological Publishing House. 240-269
[49]	Qiao T, Zhu M, 2010. Cranial morphology of the Silurian sarcopterygian Guiyu oneiros (Gnathostomata: Osteichthyes). Sci China-Earth Sci, 53(12): 1836-1848
[50]	Qiao T, Zhu M, 2015. A new Early Devonian lungfish from Guangxi, China, and its palaeogeographic significance. Alcheringa, 39(3): 428-437
[51]	Qiao T, King B, Long J A et al., 2016. Early gnathostome phylogeny revisited: multiple method consensus. PLoS One, 11(9): e0163157
[52]	Qie W K, Ma X P, Xu H H et al., 2019. Devonian integrative stratigraphy and timescale of China. Sci China-Earth Sci, 62(1): 112-134
[53]	Qu Q M, Zhu M, Wang W, 2013. Scales and dermal skeletal histology of an early bony fish Psarolepis romeri and their bearing on the evolution of rhombic scales and hard tissues. PLoS One, 8(4): e61485
[54]	Qu Q M, Haitina T, Zhu M et al., 2015. New genomic and fossil data illuminate the origin of enamel. Nature, 526: 108-111
[55]	Song C Q, Chang M M, 1991. Discovery of Chirodipterus (Dipnoi) from lower Upper Devonian of Hunan, South China. In: Chang M M, Liu Y H, Zhang G R eds. Early Vertebrates and Related Problems of Evolutionary Biology. Beijing: Science Press. 465-476
[56]	Tông-Dzuy T, Janvier P, 1990. Les Vertébrés du Dévonien inférieur du Bac Bo oriental (provinces de Bac Thaï et Lang Son, Viêt Nam). Bull Mus Natl Hist Nat, Sect C, 12(2): 143-223
[57]	Tông-Dzuy T, Janvier P, 1994. Early Devonian fishes from Trang Xa (Bac Thai, Vietnam), with remarks on the distribution of the vertebrates in the Song Cau Group. J Asian Earth Sci, 10(3/4): 235-243
[58]	Tông-Dzuy T, Phuong T H, Boucot A J et al., 1997. Silurian vertebrates from Central Vietnam (Vertébrés siluriens du Viet-nam central). C R Acad Sci, Ser IIa: Sci Terre Planet, 324(12): 1023-1030
[59]	Tông-Dzuy T, Phương T H, Janvier P et al., 2013. Silurian and Devonian in Vietnam—stratigraphy and facies. J Geodyn, 69: 165-185
[60]	Wang J Q, 2005. New material of Buchanosteus and its biostratigrapic significance. Geol Bull China, 24(9): 800-806
[61]	Wang J Q, Wang N Z, 1992. Early Devonian galeaspid Agnatha from Southeast of Yunnan, China. Vert PalAsiat, 30(3): 185-194
[62]	Wang S T, Pan J, Wang J Q, 1998. Early Devonian fishes from Central and southern Guangxi and correlation of the vertebrate biostratigraphy in South China. Vert PalAsiat, 36(1): 58-69
[63]	White E I, 1965. The head of Dipterus valenciennesi Sedgwick & Murchison. Bull Br Mus (Nat Hist), Geol, 11(1): 1-45
[64]	Xue J Z, Huang P, Wang D M et al., 2018. Silurian-Devonian terrestrial revolution in South China: taxonomy, diversity, and character evolution of vascular plants in a paleogeographically isolated, low-latitude region. Earth-Sci Rev, 180: 92-125
[65]	Yu C M, Nguyen H H, Qie W K et al., 2020. Lower Emsian biostratigraphy and event stratigraphy of Ha Giang Province, northern Vietnam. Palaeoworld, 30(1): 29-43
[66]	Yu X B, 1998. A new porolepiform-like fish, Psarolepis romeri, gen. et sp. nov. (Sarcopterygii, Osteichthyes) from the Lower Devonian of Yunnan, China. J Vert Paleont, 18(2): 261-274
[67]	Zhang S R, Shan X R, Gai Z K, 2023. New findings of huananaspid fishes from the Lower Devonian in Nanning, Guangxi and their stratigraphic and biogeographic significances. J Palaeogeogr, 25(2): 341-355
[68]	Zhao W J, Zhu M, 2010. Siluro-Devonian vertebrate biostratigraphy and biogeography of China. Palaeoworld, 19(1-2): 4-26
[69]	Zhong K, Wu Y, Yin B A et al., 1992. Stratigraphy of Guangxi, China, Part 1: Devonian of Guangxi. Wuhan: China University of Geosciences Press. 1-384
[70]	Zhu M, Ahlberg P E, 2004. The origin of the internal nostril of tetrapods. Nature, 432: 94-97
[71]	Zhu M, Fan J H, 1995. Youngolepis from the Xishancun Formation (Early Lochkovian) of Qujing, China. Geobios M S, 19: 293-299
[72]	Zhu M, Schultze H P, 2001. Interrelationships of basal osteichthyans. In: Ahlberg P E ed. Major Events in Early Vertebrate Evolution: Palaeontology, Phylogeny, Genetics and Development. London: Taylor & Francis. 289-314
[73]	Zhu M, Yu X B, 2002. A primitive fish close to the common ancestor of tetrapods and lungfish. Nature, 418: 767-770
[74]	Zhu M, Yu X B, 2004. Lower jaw character transitions among major sarcopterygian groups - a survey based on new materials from Yunnan, China. In: Arratia G, Wilson M V H, Cloutier R eds. Recent Advances in the Origin and Early Radiation of Vertebrates. München: Verlag Dr. Friedrich Pfeil. 271-286
[75]	Zhu M, Wang J Q, Fan J H, 1994. Early Devonian fishes from Guijiatun and Xujiachong formations of Qujing, Yunnan, and related biostratigraphic problems. Vert PalAsiat, 32(1): 1-20
[76]	Zhu M, Yu X B, Janvier P, 1999. A primitive fossil fish sheds light on the origin of bony fishes. Nature, 397: 607-610
[77]	Zhu M, Wang N Z, Wang J Q, 2000. Devonian macro- and microvetebrate assemblages of China. Cour Forsch Inst Senckenberg, 223: 361-372
[78]	Zhu M, Yu X B, Ahlberg P E, 2001. A primitive sarcopterygian fish with an eyestalk. Nature, 410: 81-84
[79]	Zhu M, Yu X B, Wang W et al., 2006. A primitive fish provides key characters bearing on deep osteichthyan phylogeny. Nature, 441: 77-80
[80]	Zhu M, Zhao W J, Jia L T et al., 2009. The oldest articulated osteichthyan reveals mosaic gnathostome characters. Nature, 458: 469-474
[81]	Zhu M, Yu X B, Ahlberg P E et al., 2013. A Silurian placoderm with osteichthyan-like marginal jaw bones. Nature, 502: 188-193
[82]	Zhu M, Ahlberg P E, Pan Z H et al., 2016. A Silurian maxillate placoderm illuminates jaw evolution. Science, 354: 334-336 PMID