Redescription of Nochelaspis maeandrine ,the largest eugaleaspiform from the Lower Devonian of Qujing, Yunnan
Received date: 2021-06-02
Online published: 2021-07-27
The new specimens of the largest eugaleaspiform Nochelaspis maeandrine are redescribed from two localities of the Xishancun Formation in Qujing City, Yunnan Province, southwestern China. Nochelaspis is most suggestive of Yunnanogaleaspis from the same horizon, but differs in its slit-like median dorsal opening (length/width>6), much stronger inner cornual process, coarse stellate ornamentation, and the serrations along the edges of the median dorsal opening and headshield. The new findings reveal the morphological details on the ventral side of the headshield as well. The oralobranchial fenestra is covered by a large dermal ventral plate, which is decorated with dense, tiny granular tubercles, and aligned with six pairs of separated, large, and circular branchial openings. This condition is different from that of osteostracans, in which the oralobranchial fenestra is covered by numerous minute scales or larger dermal platelets, and the branchial openings are slit-shaped and covered by small skin flaps somewhat like those of elasmobranchs. However, the branchial openings of galeaspids and osteostracans are both located ventrally as in modern rays, indicating a benthic lifestyle dwelling on sandy or muddy substrates in a quiet marine environment.
Key words： Qujing; Yunnan; Early Devonian; eugaleaspiform; Nochelaspis
MENG Xin-Yuan, ZHU Min, GAI Zhi-Kun . Redescription of Nochelaspis maeandrine ,the largest eugaleaspiform from the Lower Devonian of Qujing, Yunnan[J]. Vertebrata Palasiatica, 2021 , 59(4) : 257 -272 . DOI: 10.19615/j.cnki.2096-9899.210727
| [1] | Adam H, Straham R, 1963. Systematics and geographical distribution of myxinoids. In: Brodal A, Fänge R eds. The Biology of Myxine. Oslo: Universitetsforlaget. 1-288 |
| [2] | Bloch M E, Schneider J G, 1801. Systema Ichthyologiae: Iconibus CX Ilustratum, Vol. 2. Berolini: Sumtibus Auctoris Impressum et Bibliopolio Sanderiano Commissum. 1-584 |
| [3] | Cai C Y, Fang Z J, Li X X et al., 1994. New advance in the study of biostratigraphy of Lower and Middle Devonian marine-continental transitional strata in east Yunnan. Sci China Ser B, 24:634-639 |
| [4] | Cai J C, Zhao W J, Zhu M, 2020. Subdivision and age of the Silurian fish-bearing Kuanti Formation in Qujing, Yunnan Province. Vert PalAsiat, 58:249-266 |
| [5] | Dearden R P, Stockey C, Brazeau M D, 2019. The pharynx of the stem-chondrichthyan Ptomacanthus and the early evolution of the gnathostome gill skeleton. Nat Commun, 10:2050 |
| [6] | De Iuliis G, Pulerà D, 2007. The Dissection of Vertebrates: a Laboratory Manual. New York: Academic Press. 1-109 |
| [7] | Denison R, 1978. Placodermi. In: Schultze H P eds. Handbook of Paleoichthyology, vol. 2. Stuttgart: Gustav Fischer Verlag. 1-128 |
| [8] | Donoghue P C J, Forey P L, Aldridge R J, 2000. Conodont affinity and chordate phylogeny. Biol Rev Biol Proc Cambridge Philos Soc, 75:191-251 |
| [9] | Elliott D K, 1987. A reassessment of Astraspis desiderata , the oldest North American vertebrate. Science, 237:190-192 |
| [10] | Fernholm B, 1998. Hagfish systematics. In: Jorgensen J M, Lomholt J P, Weber R E et al. eds. The Biology of Hagfishes. London: Chapman & Hall. 33-44 |
| [11] | Forey P L, 1984. Yet more reflections on agnathan-gnathostome relationships. J Vert Palaeont, 4:330-343 |
| [12] | Gai Z K, Zhu M, 2005. A new genus of eugaleaspids (Galeaspida, Agnatha) from the Silurian of Anji, Zhejiang, China. Vert PalAsiat, 43:165-174 |
| [13] | Gai Z K, Zhu M, 2012. The origin of the vertebrate jaw: intersection between developmental biology-based model and fossil evidence. Chinese Sci Bull, 57:3819-3828 |
| [14] | Gai Z K, Zhu M, 2017. Evolutionary History of Agnathans and Their Fossil Records in China. Shanghai: Shanghai Scientific & Technical Publishers. 1-314 |
| [15] | Gai Z K, Zhu M, Zhao W J, 2005. New material of eugaleaspids from the Silurian of Changxing, Zhejiang, China, with a discussion on the eugaleaspid phylogeny. Vert PalAsiat, 43:61-75 |
| [16] | Gai Z K, Donoghue P C J, Zhu M et al., 2011. Fossil jawless fish from China foreshadows early jawed vertebrate anatomy. Nature, 476:324-327 |
| [17] | Gai Z K, Lu L W, Zhao W J, 2018. New polybranchiaspiform fishes (Agnatha: Galeaspida) from the middle Palaeozoic of China and their ecomorphological implications. PLoS One, 13:e0202217 |
| [18] | Gai Z K, Zhu M, Donoghue P C J, 2019. The circulatory system of Galeaspida (Vertebrata; stem-Gnathostomata) revealed by synchrotron X-ray tomographic microscopy. Palaeoworld, 28:441-460 |
| [19] | Gai Z K, Shan X R, Sun Z X et al., 2020. A redescription of the Silurian Sinogaleaspis shankouensis (Galeaspida, stem-Gnathostomata) from Jiangxi, China. Vert PalAsiat, 58:85-99 |
| [20] | Hughes G, 1984. General anatomy of the gills. Fish Physiol, 10:1-72 |
| [21] | Janvier P, 1996. The dawn of the vertebrates: characters versus common ascent in the rise of current vertebrate phylogenies. Palaeontology, 39:259-287 |
| [22] | Janvier P, 1996. Early Vertebrates. Oxford: Clarendon Press. 1-393 |
| [23] | Janvier P, 2007. Homologies and evolutionary transitions in early vertebrate history. In: Anderson J S, Sues H D eds. Major Transitions in Vertebrate Evolution. Bloomington and Indianapolis: Indiana University Press. 57-121 |
| [24] | Janvier P, Thanh T D, Phuong T H, 2009. Occurrence of Sanqiaspis , Liu, 1975 (Vertebrata, Galeaspida) in the Lower Devonian of Vietnam, with remarks on the anatomy and systematics of the Sanqiaspididae. C R Palevol, 8:59-65 |
| [25] | Jefferies R P S, 1986. The Ancestry of the Vertebrates. London: British Museum (Natural History). 1-376 |
| [26] | Jiang W Y, Zhu M, Shi X D et al., 2021. Qushiaspis , a new genus of gantarostrataspid fish (Galeaspida, stem-Gnathostomata) from the Lower Devonian of Yunnan, China. Hist Biol, doi: 10.1080/08912963.2021.1888086 |
| [27] | Jollie M, 1962. Chordate Morphology. New York: Reinhold Books. 1-478 |
| [28] | Kardong K V, 2015. Vertebrates: Comparative Anatomy, Function, Evolution. New York: McGraw-Hill Education. 1-795 |
| [29] | Lin B Y, Su Y Z, Zhu X F, et al. 1998. China Stratigraphic Dictionary (Silurian). Beijing: The Geological Publishing House. 1-104 |
| [30] | Liu S F, 1986. Fossil eugaleaspid from Guangxi. Vert PalAsiat, 24:1-9 |
| [31] | Liu Y H, 1965. New Devonian agnathans of Yunnan. Vert PalAsiat, 9:125-134 |
| [32] | Liu Y H, 1975. Lower Devonian agnathans of Yunnan and Sichuan. Vert PalAsiat, 13:202-216 |
| [33] | Liu Y H, 1985. A galeaspid (Agnatha), Antiquisagittaspis cornuta gen. et sp. nov., from Lower Devonian of Guangxi, China. Vert PalAsiat, 23:247-254 |
| [34] | Liu Y H, 2002. The age and correlation of some Devonian fish-bearing beds of East Yunnan, China. Vert PalAsiat, 40:52-69 |
| [35] | Liu Y H, Zhu M, Gai Z K, et al., 2015. Subclass Galeaspida. In: Zhu M ed. Palaeovertebrata Sinica, Vol. I, Fishes, Fascile 1, Agnathans. Beijing: Science Press. 141-272 |
| [36] | Liu Y H, Zhu M, Lin X H et al., 2019. A reappraisal of the Silurian galeaspids (stem-Gnathostomata) from Tarim Basin, Xinjiang. Vert PalAsiat, 57:253-273 |
| [37] | Liu Y L, Huang L B, Zong R W et al., 2020. New material of galeaspids from the Silurian Llandovery from Wuhan in South China. Earth Sci, 1-26, doi: 10.3799/dqkx.2020.343 |
| [38] | Liu Y L, Huang L B, Zong R W et al., 2021. The oldest eugaleaspiform (Galeaspida) from the Silurian Fentou Formation (Telychian, Llandovery) of Wuhan, South China. J Syst Palaeont, 19:1-16 |
| [39] | Lu L W, Pan J, Zhao L J, 2007. New findings of middle Paleozoic Agnatha and fishes from Kalpin, Xinjiang. Acta Geosci Sin, 28:143-147 |
| [40] | 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. Geol Soc London Spec Publ, 314:241-262 |
| [41] | Oisi Y, Ota K G, Fujimoto S et al., 2013. Development of the chondrocranium in hagfishes, with special reference to the early evolution of vertebrates. Zool Sci, 30:944-961 |
| [42] | Pan J, 1986a. New discovery of Silurian vertebrates in China. In: Department of Geology, Peking University ed. Selected Papers in Memory of Prof. S. H. Yoh. Beijing: Geological Publishing House. 67-75 |
| [43] | Pan J, 1986b. Note on Silurian vertebrates of China. Bull Chinese Acad Geol Sci, 15:161-190 |
| [44] | Pan J, 1992. New Galeaspids (Agnatha) from the Silurian and Devonian of China. Beijing: Geological Publishing House. 1-77 |
| [45] | Pan J, Wang S T, 1980. New finding of Galeaspiformes in South China. Acta Palaeont Sin, 19:1-7 |
| [46] | Pernègre V N, 2002. The genus Doryaspis White (Heterostraci) from the Lower Devonian of Vestspitsbergen, Svalbard. J Vert Paleont, 22:735-746 |
| [47] | Pernègre V N, 2005. Description d'une nouvelle espèce et analyse morpho-fonctionnelle du genre Doryaspis White (Heterostraci) du Dévonien du Spitsbergen. Geobios M S, 38:257-268 |
| [48] | Qie W K, Ma X P, Xu H H, 2018. Devonian integrative stratigraphy and timescale of China. Sci China Earth Sci, 62:112-134 |
| [49] | Sansom R S, Freedman K, Gabbott S E et al., 2010. Taphonomy and affinity of an enigmatic Silurian vertebrate, Jamoytius kerwoodi White. Palaeontology, 53:1393-1409 |
| [50] | Shan X R, Zhu M, Zhao W J et al., 2020. A new genus of sinogaleaspids (Galeaspida, stem-Gnathostomata) from the Silurian period in Jiangxi, China. PeerJ, 8:e9008 |
| [51] | Stensiö E A, 1968. The cyclostomes, with special reference to the diphyletic origin of the Petromyzontida and Myxinoidea. In: Ørvig T ed. Current Problems in Lower Vertebrate Phylogeny, Nobel Symposium 4. Stockholm: Almquist & Wiksell. 13-71 |
| [52] | Wang J Q, 1984. Geological and paleogeographical distribution of Devonian fishes in China. Vert PalAsiat, 22:219-229 |
| [53] | Wang N Z, 1991. Two new Silurian galeaspids (jawless craniates) from Zhejiang Province, China, with a discussion of galeaspid-gnathostome relationships. In: Chang M M, Liu Y H, Zhang G R eds. Early Vertebrates and Related Problems of Evolutionary Biology. Beijing: Science Press. 41-66 |
| [54] | Xue J Z, Huang P, Wang D M, 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 |
| [55] | Zhao W J, Zhu M, 2010. Siluro-Devonian vertebrate biostratigraphy and biogeography of China. Palaeoworld, 19:4-26 |
| [56] | Zhu M, 1992. Two new eugaleaspids, with a discussion on eugaleaspid phylogeny. Vert PalAsiat, 30:169-184 |
| [57] | Zhu M, 2000. Catalogue of Devonian vertebrates in China, with notes on bio-events. Courier Forschungsinst Senckenberg, 223:373-390 |
| [58] | Zhu M, Gai Z K, 2006. Phylogenetic relationships of galeaspids (Agnatha). Vert PalAsiat, 44:1-27 |
| [59] | Zhu M, Liu Y H, Jia L T et al., 2012. A new genus of eugaleaspidiforms (Agnatha: Galeaspida) from the Ludlow, Silurian of Qujing, Yunnan, southwestern China. Vert PalAsiat, 50:1-7 |
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