Reappraisal of Bothriolepis sinensis  Chi, 1940 from the Tiaomachien Formation, Hunan, China

doi:10.19615/j.cnki.2096-9899.230901

Abstract

Abstract:

Bothriolepis sinensis Chi 1940, mainly based on anterior median dorsal plates from the Middle Devonian Tiaomachien Formation of Hunan, is the first Paleozoic vertebrate taxon erected in China. Although additional materials of B. sinensis from the type locality were described by Lu in 1988, its morphology and phylogeny remain poorly understood. In this study, we complemented the morphology of the skull and trunk armor of B. sinensis based on Chi’s specimens housed in the Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, and several previously undescribed specimens in the Geological Museum of China. Bothriolepis sinensis differs from other Bothriolepis in the following combination of characteristics: enlarged supraotic thickening, length/width ratio of head shield 1.4-1.6, broad orbital fenestra (greater than 1/3 of the head shield width), and fan-shaped preorbital recess. The phylogenetic analysis did not place B. askinae in the most basal position of the genus and revealed that B. sinensis and B. kwangtungensis consistently from a monophyletic group characterized by their slender proximal segment of the pectoral fin (length/width ratio greater than 7). A majority of Chinese Bothriolepis species (B. niushoushanensis , B. lochangensis , B. tungseni , B. kwangtungensis and B. sinensis ) were clustered in a clade characterized by the pectoral pit-line on the ventral central plate 1 extending to the ventral central plate 2. The paleogeographic reconstruction using the data from the DeepBone platform showed that Bothriolepis had its oldest occurrences in South China and East Gondwana in Eifelian, dispersed rapidly worldwide, and then diversified across the coasts of the Rheic Ocean.

Key words: Hunan, Middle Devonian, Antiarcha, Bothriolepis , phylogeny, paleobiogeography, data visualization

摘要：

计荣森先生于1940年基于湖南长沙附近中泥盆统跳马涧组采集到的一些以前中背片为代表的胴甲鱼类标本命名了中华沟鳞鱼(Bothriolepis sinensis ), 这是在中国正式命名的第一种古生代脊椎动物。虽然卢立伍于1988年对后续出产于模式种产地的其他中华沟鳞鱼材料进行了补充描述，但研究者对中华沟鳞鱼的形态学特征和系统发育仍缺乏全面认识。在此基于计荣森先生所采集的样本及一些收藏于中国地质博物馆未正式发表的化石，对中华沟鳞鱼进行了再研究，补充了其头部与躯甲的部分特征。与其他沟鳞鱼类相比，中华沟鳞鱼拥有以下一系列特征：较大的听上加厚区、颅顶甲长宽比1.4-1.6、较宽的眶窗(宽于颅顶甲的1/3)以及扇形的眶前凹等。系统发育分析结果显示，B. askinae 并非位于沟鳞鱼支系的最原始位置；中华沟鳞鱼与广东沟鳞鱼因同样拥有纤细的附肢近端(长宽比大于7)而形成了一个稳定的单系。本研究中绝大部分的中国种类均位于同一支系上，该支系的共有衍征为其胸鳍感觉管从第一腹中片延伸至第二腹中片。此外，基于“深骨”平台数据的古生物地理可视化分析结果表明，沟鳞鱼类最早于艾菲尔期出现在华南与东冈瓦纳，并迅速地辐射并扩散至全球，最终于法门期在瑞亚克洋两岸表现出了高度的物种多样性。

关键词: 湖南，中泥盆统，胴甲鱼类，沟鳞鱼类，系统发育，古生物地理学，数据可视化

CLC Number:

Q915.862

LUO Yan-Chao, ZHU Min, LU Li-Wu, PAN Zhao-Hui. Reappraisal of Bothriolepis sinensis Chi, 1940 from the Tiaomachien Formation, Hunan, China. Vertebrata Palasiatica, 2023, 61(4): 261-276.

罗彦超, 朱敏, 卢立伍, 潘照晖. 2023, 61(4): 261-276, 湖南中泥盆统跳马涧组中华沟鳞鱼再研究. 古脊椎动物学报.

Figures/Tables 7

Fig. 1 Map of the fossil site and the fish-bearing horizon in this study Based on: GS(2019)3266

Fig. 2 Photo (A) and illustrative drawing (B) of the referred specimen of Bothriolepis sinensis (GMV1961)

Fig. 3 Anterior median dorsal plate of Bothriolepis sinensis A. photo of the internal mould of AMD (IVPP V467a, Chi, 1940:pl. I-1); B, C. photo and illustrative drawing of AMD (V467a) in ventral view; D. internal mould of AMD (V467c, Chi, 1940:pl. I-3) A and D are coated with ammonium chloride. Scale bars = 10 mm

Fig. 4 Other referred specimens of B. sinensis collected by Chi in the 1930s A. internal mould of the left ADL (IVPP V467b, Chi, 1940:pl. I-7); B. internal mould of the right MxL (V467d, Chi, 1940:pl. I-5). Fossils are whitened with ammonium chloride Scale bars = 10 mm

Fig. 5 Reconstruction of the trunk armour of Bothriolepis sinensis

Fig. 6 The 50% majority rule consensus tree of 264 most parsimonious trees, shows the phylogenetic position of B. sinensis within Bothriolepis Numbers on the branches display the majorities which do not reach 100

Fig. 7 Geographical distribution of Bothriolepis A. fossil sites of Bothriolepis ; B. paleobiogeography of Bothriolepis based on Scotese’s paleogeographic reconstructions. Abbreviations: Aus, Australia; Bal. Baltica; Lau. Laurentia; SC. South China; Sib. Siberia

References 67

[1]	Brazeau M D, 2009. The braincase and jaws of a Devonian ‘acanthodian’ and modern gnathostome origins. Nature, 457: 305-308 DOI
[2]	Carr R K, 1995. Placoderm diversity and evolution. Bull Mus Natl Hist Nat, Paris, Sér 4, Sect C, 17: 85-125
[3]	Chang K J, 1965. New antiarchs from the Middle Devonian of Yunnan. Vert PalAsiat, 9: 1-9
[4]	Charest F, Johanson Z, Cloutier R, 2018. Loss in the making: absence of pelvic fins and presence of paedomorphic pelvic girdles in a Late Devonian antiarch placoderm (jawed stem-gnathostome). Biol Lett, 14: 20180199 DOI URL
[5]	Chi Y S, 1940. On the discovery of Bothriolepis in the Devonian of central Hunan. Bull Geol Soc China, 20: 57-73 DOI URL
[6]	Cope E D, 1885. The position of Pterichthys in the system. Am Nat, 19: 289-291 DOI URL
[7]	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 DOI
[8]	Downs J P, Daeschler E B, Garcia V E et al., 2016. A new large-bodied species of Bothriolepis (Antiarchi) from the Upper Devonian of Ellesmere Island, Nunavut, Canada. J Vert Paleont, 36: e1221833 DOI URL
[9]	Dupret V, Sanchez S, Goujet D et al., 2014. A primitive placoderm sheds light on the origin of the jawed vertebrate face. Nature, 507: 500-503 DOI
[10]	Dupret V, Byrne H M, Castro N et al., 2023. The Bothriolepis (Placodermi, Antiarcha) material from the Valentia Slate Formation of the Iveragh Peninsula (middle Givetian, Ireland): morphology, evolutionary and systematic considerations, phylogenetic and palaeogeographic implications. PLoS One, 18: e0280208 DOI URL
[11]	Eichwald E, 1840. Die Tier-und Pflanzenreste des alten roten Sandsteins und Bergkalks im Nowgorodschen Gouvernement. Saint-Petersburg Acad Imp Sci Bull Sci, 7: 78-91
[12]	Giles S, Rücklin M, Donoghue P C J, 2013. Histology of “placoderm” dermal skeletons: implications for the nature of the ancestral gnathostome. J Morphol, 274: 627-644 DOI PMID
[13]	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 DOI URL
[14]	Goloboff P A, Catalano S A, 2016. TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics, 32: 221-238 DOI PMID
[15]	Goujet D F, Young G C, 1995. Interrelationships of placoderms revisited. Geobios Mém Spéc, 19: 89-95
[16]	Gross W, 1965. Bothriolepis cf. panderi Lahusen in einem Geschiebe von Travemünde bei Lübeck. Mitt Geol Staatsinst Hamburg, 34: 138-141
[17]	Janvier P, 1996. The dawn of the vertebrates: characters versus common ascent in the rise of current vertebrate phylogenies. Palaeontology, 39: 259-287
[18]	Janvier P, Pan J, 1982. Hyrcanaspis bliecki n. g. n. sp., a new primitive euantiarch (Antiarcha, Placodermi) from the Middle Devonian of northeastern Iran, with a discussion on antiarch phylogeny. N J Geol Palaont Abh, 164: 364-392
[19]	Johanson Z, 1997. New antiarchs (Placodermi) from the Hunter Siltstone (Famennian) near Grenfell, N.S.W. Alcheringa, 21: 191-217 DOI URL
[20]	Ke Y, Shen S Z, Shi G R et al., 2016. Global brachiopod palaeobiogeographical evolution from Changhsingian (Late Permian) to Rhaetian (Late Triassic). Palaeogeogr, Palaeoclimatol, Palaeoecol, 448: 4-25 DOI URL
[21]	Leidy J, 1856. Descriptions of some remains of fishes from the Carboniferous and Devonian formations of the United States. J Acad Nat Sci Phila, 3: 159-165
[22]	Liu Y H, 1962. A new species of Bothriolepis from Yunnan. Vert PalAsiat, 6: 80-85
[23]	Long J A, 1983. New bothriolepid fish from the Late Devonian of Victoria, Australia. Palaeontology, 26: 295-320
[24]	Long J A, Werdelin L, 1986. A new Late Devonian bothriolepid (Placodermi, Antiarcha) from Victoria, with descriptions of other species from the state. Alcheringa, 10: 355-399 DOI URL
[25]	Long J A, Mark-Kurik E, Johanson Z et al., 2015. Copulation in antiarch placoderms and the origin of gnathostome internal fertilization. Nature, 517: 196-199 DOI
[26]	Lu L W, 1988. New data of Bothriolepis (Placodermi:Antiaroha) from Middle Devonian of Central Hunan, China. In: Xin Y S, Huang Z Z, Wang F Q et al. eds. Geological Museum Study. Beijing: Seismological Press. 69-77
[27]	Lukševičs E, 1996. Bothriolepid antiarchs (Placodermi, Bothriolepididae) from the North-western part of East European platform. Ph. D thesis. Riga: University of Latvia. Summary
[28]	Lukševičs E, 2001. Bothriolepid antiarchs (Vertebrata, Placodermi) from the Devonian of the north-western part of the East European Platform. Geodiversitas, 23: 489-609
[29]	Lukševičs E, 2021. Revision of asterolepidoid antiarch remains from the Ogre Formation (Upper Devonian) of Latvia. Proc Est Acad Sci Geol, 70: 3-17
[30]	Maddison W P, Maddison D R, 2019. Mesquite: a modular system for evolutionary analysis. Version 3.61. Updated at http://www.mesquiteproject.org, accessed 21 February 2021
[31]	McCoy F, 1848. On some new fossil fish from the Carboniferous Period. Ann Mag Nat Hist, 2: 1-10, 115-133
[32]	Miles R S, 1968. The Old Red Sandstone antiarchs of Scotland: Family Bothriolepididae. Palaeontogr Soc Monogr, 122: 1-130
[33]	Miller H, 1841. The Old Red Sandstone or New Walks in an Old Field. Edinburgh: John Johnstone. 1-275
[34]	Moloshnikov S V, 2004. Crested antiarch Bothriolepis zadonica HD Obrucheva from the Lower Famennian of Central European Russia. Acta Palaeontol Pol, 49: 135-146
[35]	Moloshnikov S V, 2008. Devonian antiarchs (Pisces, Antiarchi) from central and Southern European Russia. Paleontol J, 42: 691-773 DOI URL
[36]	Moloshnikov S V, 2009. New data on Late Devonian bothriolepidid placoderms (Pisces, Antiarchi) from Tuva. Paleontol J, 43: 558-568 DOI URL
[37]	Moy-Thomas J A, Miles R S, 1971. Palaeozoic Fishes. London: Chapman and Hall. 1-259
[38]	Pan J, 1958. On the age of Tiaomachien series of Hunan. Acta Geol Sin, 38: 135-144
[39]	Pan J, 1959. Devonian fish fossils of China and their stratigraphic and geographic distributions. Monogr Sum Basic Data Chinese Geol, 1: 23-34
[40]	Pan J, Dineley D L, 1988. A review of early (Silurian and Devonian) vertebrate biogeography and biostratigraphy of China. Proc R Soc B, 235: 29-61
[41]	Pan J, Wang S T, Liu S Y et al., 1980. Discovery of Devonian Bothriolepis and Remigolepis in Ningxia. Acta Geol Sin, 3: 175-185
[42]	Pan J, Huo F C, Cao J X et al., 1987. Continental Devonian System of Ningxia and its Biotas. Beijing: Geological Publishing House. 1-237
[43]	Pan Z H, Zhu M, 2019. VPPDB: Hosting the data for Vertebrate Paleoanthropology. Acta Geol Sin Engl Ed, 93: 61-63
[44]	Pan Z H, Zhu M, Zhu Y A et al., 2018. A new antiarch placoderm from the Emsian (Early Devonian) of Wuding, Yunnan, China. Alcheringa, 17: 1-19 DOI URL
[45]	Pan Z H, Niu Z B, Xian Z M et al., 2022. A novel specimen-based mid-Paleozoic dataset of antiarch placoderms (the most basal jawed vertebrates). Earth Syst Sci Data, 15: 41-51 DOI URL
[46]	Qiao T, Zhu M, 2015. A new Early Devonian lungfish from Guangxi, China, and its palaeogeographic significance. Alcheringa, 39: 428-437 DOI URL
[47]	Qiao T, King B, Long J A et al., 2016. Early gnathostome phylogeny revisited: multiple method consensus. PLoS One, 11: e0163157 DOI URL
[48]	Qie W K, Liang K, Königshof P, 2019. Devonian palaeoecosystems and palaeoenvironments of South China. Paleobio Paleoenv, 99: 1-5
[49]	Scotese C R, 2002. Palaeo Map Project. Available at www.scotese.com
[50]	Scotese C R, Song H, Mills B J W et al., 2021. Phanerozoic paleotemperatures: the earth’s changing climate during the last 540 million years. Earth Sci Rev, 215: 103503 DOI URL
[51]	Stensiö E A, 1948. On the Placodermi of the Upper Devonian of East Greenland. II. Antiarchi: subfamily Bothriolepinae. With an attempt at a revision of the previously described species of that family. Medd Grønl, 139: 1-622
[52]	Thomson K S, Thomas B, 2001. On the status of species of Bothriolepis (Placodermi, Antiarchi) in North America. J Vert Paleont, 21: 679-686 DOI URL
[53]	Trinajstic K, Boisvert C, Long J A et al., 2015. Pelvic and reproductive structures in placoderms (stem gnathostomes). Biol Rev, 90: 467-501 DOI URL
[54]	Wang G T, 1984. The investigation of sedimentary facies of Tiaomajian Formation in Tiaomajian Town of Changsha Country. Hunan Geol, 3: 45-56, 72
[55]	Wang J Q, 1991. The Antiarchi from Early Silurian of Hunan. Vert PalAsiat, 29: 240-244
[56]	Wang Y J, Zhu M, 2018. Redescription of Phymolepis cuifengshanensis (Antiarcha: Yunnanolepididae) using high-resolution computed tomography and new insights into anatomical details of the endocranium in antiarchs. PeerJ, 6: e4808 DOI URL
[57]	Wang Y J, Zhu M, 2021. New data on the headshield of Parayunnanolepis xitunensis (Placodermi, Antiarcha), with comments on nasal capsules in antiarchs. J Vert Paleont, 40: e1855189 DOI URL
[58]	Young G C, 1988. Antiarchs (placoderm fishes) from the Devonian Aztec Siltstone, Southern Victoria Land, Antarctica. Palaeontogr Abt A, 202: 1-125
[59]	Young G C, Friedrich Pfeil. 2010. A new antiarch (placoderm fish:Devonian) from the south coast of New South Wales, Australia. In: Elliott D K, Maisey J G, Yu X et al. eds. Morphology, Phylogeny and Paleobiogeography of Fossil Fishes (Honoring Meemann Chang). Munich: Verlag Dr. 85-100
[60]	Young G C, Gorter J D, 1981. A new fish fauna of Middle Devonian age from the Taemas/Wee Jasper region of New South Wales. Bull Bur Min Res Geol Geophys, 209: 85-147
[61]	Zhao W J, Zhu M, 2010. Siluro-Devonian vertebrate biostratigraphy and biogeography of China. Palaeoworld, 19: 4-26 DOI URL
[62]	Zhao W J, Zhu M, Liu S et al., 2016. A new look at the Silurian fish-bearing strata around the Shanmen Reservoir in Lixian, Hunan Province. J Stratigr, 40: 341-350
[63]	Zhu M, Yu X B, Choo B et al., 2012. An antiarch placoderm shows that pelvic girdles arose at the root of jawed vertebrates. Biol Lett, 8: 453-456 DOI URL
[64]	Zhu M, Yu X B, Ahlberg P E et al., 2013. A Silurian placoderm with osteichthyan-like marginal jaw bones. Nature, 502: 188-193 DOI
[65]	Zhu M, Ahlberg P E, Pan Z H et al., 2016. A Silurian maxillate placoderm illuminates jaw evolution. Science, 354: 34-336 DOI URL
[66]	Zhu Y A, Giles S, Young G C et al., 2021. Endocast and bony labyrinth of a Devonian “placoderm” challenges stem gnathostome phylogeny. Curr Biol, 31: 1112-1118 DOI URL
[67]	Zhu Y A, Li Q, Lu J et al., 2022. The oldest complete jawed vertebrates from the early Silurian of China. Nature, 609: 954-958 DOI