New findings of huananaspid fishes from the Lower Devonian in Nanning, Guangxi and their stratigraphic and biogeographic significances
ZHANG Shurong1,2, SHAN Xianren1,2, GAI Zhikun1,2
1 Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences,Beijing 100044,China; 2 University of Chinese Academy of Sciences,Beijing 100049,China
Abstract：Two new species of Huananaspidae(Huananaspidiformes,Galeaspida),Huananaspis cupido sp. nov. and Asiaspis liangi sp. nov.,are described from the lower part of the Nagaoling Formation(Pragian,Early Devonian)in Nanning,Guangxi,China. H. cupido sp. nov. differs from H. wudingensis in smaller headshield,lacking median dorsal spine,and cornual processes extending beyond the posterior margin of the headshield. A. liangi sp. nov. differs from A. expansa in headshield length greater than width,cornual processes extending posterolaterally,serrated posterior margin of cornual processes,and longitudinal oval median dorsal opening. The Nagaoling Formation yields reliable index fossils indicating a Pragian age,thus the Xujiachong vertebrate assemblage yielded from this formation can provide a palaeoichthyological standard for stratigraphic correlations of Early Devonian in South China. New findings of Huananaspis and Asiaspis from the Nagaoling Formation enriches the biodiversity of huananaspids,adds new members to the the Xujiachong vertebrate assemblage,and provides direct fossil evidence for the correlation between the Posongchong Formation in Wuding of Yunnan and the Nagaoling Formation in Nanning of Guangxi. The Xujiachong vertebrate assemblage is the most diversified in the transitional beds between the Lianhuashan and Nagaoling formations,almost containing all genera in adjacent regions,indicating that Naning of Guangxi is one of the centers for galeaspids radiating and dispersing. Fish assemblage displays a similar high level of endemism to that of the vascular plants and brachiopod fauna during the Pragian of Early Devonian,which probably caused by the South China sea,a large semi-closed epicontinental gulf. Because of the effect of a closed sea,fishes show exceptional taxonomic diversity and high levels of endemism,forming an independent biogeographic region known as‘galeaspid-yunnanolepid province’ in South China.
ZHANG Shurong,SHAN Xianren,GAI Zhikun. New findings of huananaspid fishes from the Lower Devonian in Nanning, Guangxi and their stratigraphic and biogeographic significances[J]. JOPC, 2023, 25(2): 341-355.
[1] 盖志琨,朱敏. 2007. 云南早泥盆世西山村组华南鱼类的首次发现. 古脊椎动物学报, 45(1): 1-12. [Gai Z K,Zhu M.2007. First discovery of Huananaspidae from the Xishancun formation(Lochkovian,Devonian)of Yunnan,China. Vertebrata PalAsiatica, 45(1): 1-12] [2] 郝守刚. 1988. 早泥盆世始叶蕨(新属)及叶子起源的讨论. 植物学报, 30(4): 441-448. [Hao S G.1988. A new Lower Devonian genus from Yunnan,with notes on the origin of leaf. Journal of Integrative Plant Biology, 30(4): 441-448] [3] 姬书安,潘江. 1999. 滇东泥盆纪瓣甲鱼类(盾皮鱼纲)化石新知. 地层古生物论文集, 27: 172-179. [Ji S A,Pan J.1999. New Devonian petalichthyids(Placodermi)from eastern Yunnan,China. Professional Papers of Stratigraphy and Palaeontology, 27: 172-179] [4] 邝国敦,赵明特,陶业斌. 1989. 中国海相泥盆系标准剖面: 广西六景泥盆系剖面. 武汉: 中国地质大学出版社,1-145. [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,1-145] [5] 廖卫华,许汉奎,王成源,阮亦萍,蔡重阳,穆道成,卢礼昌. 1978. 西南地区泥盆纪地层的划分与对比. 见: 南京地质古生物研究所. 西南地区碳酸盐生物地层. 北京: 地质出版社,221-249. [Liao W H,Xu H K,Wang C Y,Ruan Y P,Cai C Y,Mu D C,Lu L C.1978. On some basic Devonian sections in Southwestern China. In: Nanjing Institute of Geology and Palaeontology. Carbonate Biostratigraphy in Southwestern Regions. Beijing: Science Press,221-249] [6] 刘时藩. 1982a. 广西六景节甲鱼化石. 古脊椎动物与古人类, 20(2): 106-114. [Liu S F.1982a. Preliminary note on the Arthrodira from Guangxi,China. Vertebrata PalAsiatica, 20(2): 106-114] [7] 刘时藩. 1982b. 节甲鱼的一内颅化石. 古脊椎动物学报, 20(4): 272-275. [Liu S F.1982b. An arthrodiren endocranium. Vertebrata PalAsiatica, 20(4): 272-275] [8] 刘时藩. 1986. 广西盔甲鱼类化石. 古脊椎动物学报, 24(1): 1-9. [Liu S F.1986. Fossil eugaleaspid from Guangxi. Vertebrata PalAsiatica, 24(1): 1-9] [9] 刘时藩. 1992. 桂中南下泥盆统的胴甲鱼类化石. 古脊椎动物学报, 30(3): 210-220. [Liu S F.1992. On the Lower Devonian antiarchians of Guangxi,China. Vertebrata PalAsiatica, 30(3): 210-220] [10] 刘时藩. 1994. 滇东武定地区的中泥盆统. 古脊椎动物学报, 32(1): 32-40. [Liu S F.1994. On the Middle Devonian series of Wuding,Yunnan. Vertebrata PalAsiatica, 32(1): 32-40] [11] 刘玉海. 1965. 云南曲靖地区早泥盆世无颌类化石. 古脊椎动物学报, 9(2): 125-134. [Liu Y H.1965. New Devonian agnathans of Yunnan. Vertebrata PalAsiatica, 9(2): 125-134] [12] 刘玉海. 1973. 川滇泥盆纪的多鳃鱼和大瓣鱼化石. 古脊椎动物与古人类, 11(2): 132-143. [Liu Y H.1973. On the new forms of Polybranchiaspiformes and Petalichthyida from Devonian of Southwest China. Vertebrata PalAsiatica, 11(2): 132-143] [13] 刘玉海. 1975. 川滇早泥盆世的无颌类. 古脊椎动物学报, 13(4): 202-216. [Liu Y H.1975. Lower Devonian agnathans of Yunnan and Sichuan. Vertebrata PalAsiatica, 13(4): 202-216] [14] 刘玉海. 1985. 盔甲鱼类Antiquisagittaspis cornuta(新属、新种)在广西六景下泥盆统的发现. 古脊椎动物学报, 23(4): 247-254,317. [Liu Y H.1985. A galeaspid(Agnatha),Antiquasagittaspis cornuta gen. et sp. nov.,from Lower Devonian of Guangxi,China. Vertebrata PalAsiatica, 23(4): 247-254,317] [15] 潘江,王士涛,刘运鹏. 1975. 中国南方早泥盆世无颌类及鱼类化石. 地层古生物论文集, 1: 135-169,285-301,315. [P’an K,Wang S T,Liu Y P.1975. The Lower Devonian agnatha and pisces from South China. Professional Papers of Stratigraphy and Paleontology, 1: 135-169,285-301,315] [16] 潘江,王士涛. 1978. 中国南方泥盆纪无颌类及鱼类化石. 见: 中国地质科学院地质与矿产资源研究所. 华南泥盆系会议论文集. 北京: 地质出版社,298-333. [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 Sciences. Symposium on the Devonian System of South China. Beijing: Geological Publishing House,298-333] [17] 潘江,王士涛. 1981. 云南早泥盆世多鳃鱼类的新发现. 古脊椎动物学报, 19(2): 113-121. [Pan J,Wang S T.1981. New discoveries of polybranchiaspids from Yunnan Province. Vertebrata PalAsiatica, 19(2): 113-121] [18] 王俊卿,王念忠. 1992. 滇东南早泥盆世无颌类. 古脊椎动物学报, 30(3): 185-194. [Wang J Q,Wang N Z.1992. Early Devonian galeaspid agnatha from Southeast of Yunnan,China. Vertebrata PalAsiatica, 30(3): 185-194] [19] 王俊卿,朱敏. 1994. 滇东北昭通早泥盆世盔甲鱼类一新属种. 古脊椎动物学报, 32(4): 231-243. [Wang J Q,Zhu M.1994. Zhaotongaspis janvieri gen. et sp. nov.,a galeaspid from Early Devonian of Zhaotong,northeastern Yunnan. Vertebrata PalAsiatica, 32(4): 231-243] [20] 王俊卿,范俊航,朱敏. 1996. 滇东北昭通附近早泥盆世脊椎动物化石新知. 古脊椎动物学报, 34(1): 1-17. [Wang J Q,Fan J H,Zhu M.1996. Early vertebrate fossils from the Early Devonian of Zhaotong District,northeastern Yunnan. Vertebrata PalAsiatica, 34(1): 1-17] [21] 王俊卿,盖志琨,朱敏. 2005. 云南文山大窗鱼属(Macrothyraspis)一新种. 古脊椎动物学报, 43(4): 304-311. [Wang J Q,Gai Z K,Zhu M.2005. A new species of Macrothyraspis(Galeaspida,Agnatha)from Wenshan,Yunnan,China. Vertebrata PalAsiatica, 43(4): 304-311] [22] 王士涛,潘江,王俊卿. 1998. 广西早泥盆世节甲鱼类及胴甲鱼类化石及华南早泥盆世脊椎动物生物地层对比. 古脊椎动物学报, 36(1): 58-69. [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. Vertebrata PalAsiatica, 36(1): 58-69] [23] 王怿. 1994. 滇东南文山古木早泥盆世孢子组合. 微体古生物学报, 11(3): 319-332. [Wang Y.1994. Lower Devonian miospores from Gumu in the Wenshan district,southeastern Yunnan. Acta Micropalaeontologica Sinica, 11(3): 319-332] [24] 王钰,俞昌民,方大卫. 1964. 广西中东部泥盆系那高岭组的时代及郁江组的分层. 科学通报, 15(11): 1013-1016. [Wang Y,Yu C M,Fang D W.1964. Age of the Nagaoling Formation and division of the Yujiang Formation in central eastern Guangxi Province. Science Bulletin, 15(11): 1013-1016] [25] 曾广春,纵瑞文,刘琦. 2018. 隐藏的风景: 广西古生物化石记. 南宁: 广西美术出版社,1-511. [Zeng G C,Zong R W,Liu Q.2018. Hidden Scenery: Fossil Hunting in Guangxi. Nanning: Guangxi Fine Arts Publishing House,1-511] [26] 钟铿,吴诒,殷保安,梁演林,姚肇贵,彭金兰. 1992. 广西的泥盆系: 广西地层之一. 武汉: 中国地质大学出版社,1-384. [Zhong K,Wu Y,Yin B A,Liang Y L,Yao Z G,Peng J L.1992. Stratigraphy of Guangxi,China,Part 1: Devonian of Guangxi. Wuhan: China University of Geosciences Press,1-384] [27] 赵文金,朱敏,贾连涛. 2002. 云南文山早泥盆世盔甲鱼类的新发现. 古脊椎动物学报, 40(2): 97-113. [Zhao W J,Zhu M,Jia L T.2002. New discovery of galeaspids from Early Devonian of Wenshan,southeastern Yunnan,China. Vertebrata PalAsiatica, 40(2): 97-113] [28] 朱敏,王俊卿,范俊航. 1994. 云南曲靖地区桂家屯组与徐家冲组早期脊椎动物化石及相关生物地层问题. 古脊椎动物学报, 32(1): 1-20. [Zhu M,Wang J Q,Fan J H.1994. Early Devonian fishes from Guijiatun and Xujiachong Formations of Qujing,Yunnan,and related biostratigraphic problems. Vertebrata PalAsiatica, 32(1): 1-20] [29] Dupret V.2008. A new wuttagoonaspid(Placodermi,Arthrodira)from the Lower Devonian of Yunnan(South China): origin,dispersal,and paleobiogeographic significance. Journal of Vertebrate Paleontology, 28(1): 12-20. [30] 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. Zoological Journal of the Linnean Society, 157(1): 70-82. [31] Gai Z K,Zhu M,Jia L T,Zhao W J.2015. A streamlined jawless fish(Galeaspida)from the Lower Devonian of Yunnan,China and its taxonomic and paleoecological implications. Vertebrata PalAsiatica, 53(2): 93-109. [32] Gai Z K, Lu L W, Zhao W J, Zhu M.2018. New polybranchiaspiform fishes (Agnatha: Galeaspida) from the Middle Palaeozoic of China and their ecomorphological implications. Plos One, 13: e0202217. [33] Gai Z K,Jiang W Y,Zhao W J,Li Q,Shi X D,Zhu M.2022. Redescription of the Sanqiaspidae(Galeaspida)from the Lower Devonian of South China and its biostratigraphic significance. Palaeobiodiversity and Palaeoenvironments, 102(1): 173-191. [34] 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. Palaeobiodiversity and Palaeoenvironments, 99(1): 29-43. [35] Huang B C,Yan Y G,Piper J D,Zhang D H,Yi Z Y,Yu S,Zhou T H.2018. Paleomagnetic constraints on the paleogeography of the East Asian blocks during Late Paleozoic and Early Mesozoic times. Earth-Science Reviews, 186: 8-36. [36] Janvier P.1996. Early Vertebrates. Oxford: Clarendon Press,1-393. [37] Janvier P,Thanh T D,Phuong T H,Clément G,Phong N D.2009. Occurrence of Sanqiaspis,Liu,1975(Vertebrata,Galeaspida)in the Lower Devonian of Vietnam,with remarks on the anatomy and systematics of the Sanqiaspididae. Comptes Rendus Palevol, 8(1): 59-65. [38] 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. Geological Society,Special Publications,London, 314(1): 241-262. [39] Meng X Y,Zhu M,Wang J Q,Pan Z H,Gai Z K.2022. First Middle Devonian galeaspid from the Haikou Formation in Yunnan Province. Vertebrata PalAsiatica, 60(3): 184-196. [40] Lu J,Zhu M.2008. An Early Devonian(Pragian)sarcopterygian from Zhaotong,Yunnan,China. Vertebrata PalAsiatica, 46(3): 161-170. [41] Lu J,Zhu M.2010. An onychodont fish(Osteichthyes,Sarcopterygii)from the Early Devonian of China,and the evolution of the Onychodontiformes. Proceedings of the Royal Society B: Biological Sciences, 277(1679): 293-299. [42] Lu J,Zhu M,Long J A,Zhao W,Senden T J,Jia L,Qiao T.2012. The earliest known stem-tetrapod from the Lower Devonian of China. Nature Communications, 3: 1160. [43] Lu J F,Qie W K,Chen X Q.2016. Pragian and lower Emsian(Lower Devonian)conodonts from Liujing,Guangxi,South China. Alcheringa, 40(2): 275-296. [44] Pan J.1992. New Galeaspids(Agnatha)from the Silurian and Devonian of China. Beijing: Geological Publishing House,1-77. [45] Qiao T,Zhu M.2015. A new Early Devonian lungfish from Guangxi,China,and its palaeogeographic significance. Alcheringa, 39(3): 428-437. [46] Qie W K,Ma X P,Xu H H,Qiao L,Liang K,Guo W,Song J J,Chen B,Lu J F.2019. Devonian integrative stratigraphy and timescale of China. Science China: Earth Sciences, 62(1): 112-134. [47] Tarlo L B H. 1967. Agnatha. In: Harland W B,Holland C H,House M R,Hughes N F,Reynolds A B,Rudwick,M J S,Satterthwaite G E,Tarlo L B H,Willey E C(eds). The Fossil Record: a symposium with documentation. London: The Geological Society of London,629-636. [48] Tông Dzuy T,Janvier P,Phuong T H.1996. Fish suggests continental connections between the IndoChina and South China blocks in Middle Devonian time. Geology, 24(6): 571-574. [49] Torsvik T H,Cocks L R M.2004. Earth geography from 400 to 250 Ma: a palaeomagnetic,faunal and facies review. Journal of the Geological Society,London, 161: 555-572. [50] Xue J Z,Huang P,Wang D M,Xiong C H,Liu L,Basinger J F.2018. Silurian-Devonian terrestrial revolution in South China: taxonomy,diversity,and character evolution of vascular plants in a paleogeographically isolated,low-latitude region. Earth-Science Reviews, 180: 92-125. [51] Young G C.1981. Biogeography of Devonian vertebrates. Alcheringa, 5(3): 225-243. [52] Young G C.1993. Vertebrate faunal provinces in the Middle Palaeozoic. In: Long J A. Palaeozoic Vertebrate Biostratigraphy and Biogeography. London: Belhaven Press,293-323. [53] Young G C,Janvier P.1999. Early-Middle Palaeozoic vertebrate faunas in relation to Gondwana dispersion and Asian accretion. In: Metcalfe I,Ren J S,Charvet J,Hada S. Gondwana Dispersion and Asian Accretion. IGCP 321 Final Results Volume. Rotterdam: A. A. Balkema,115-140. [54] Yu C M,Qie W K,Lu J F.2018. Emsian(Early Devonian)Yujiang Event in South China. Palaeoworld, 27(1): 53-65. [55] Zhang G W,Guo A L,Wang Y J,Li S Z,Dong Y P,Liu S F,He D F,Cheng S,Lu R K,Yao A P.2013. Tectonics of South China continent and its implications. Science China Earth Sciences, 56(11): 1804-1828. [56] Zhang M Q,Ma X P.2019. Origination and diversification of Devonian ambocoelioid brachiopods in South China. Palaeobiodiversity and Palaeoenvironments, 99(1): 63-90. [57] Zhao W J,Zhu M.2010. Siluro-Devonian vertebrate biostratigraphy and biogeography of China. Palaeoworld, 19(1-2): 4-26. [58] Zhu M,Wang N Z,Wang J Q.2000. Devonian macro- and microvetebrate assemblages of China. Courier Forschungsinstitut Senckenberg, 223: 361-372. [59] Zhu M,Gai Z K.2006. Phylogenetic relationships of galeaspids(Agnatha). Vertebrata PalAsiatica, 44(1): 1-27. [60] Janvier P.1975. Anatomie et position systematique des Galeaspides(Vertebrata,Cyclostomata),Cephalaspidomorphes du Devonien inferieur du Yunnan(Chine). Muséum National d’Histoire Naturelle,Paris, 278: 1-16.
