A Middle Triassic kyphosichthyiform from Yunnan, China, and phylogenetic reassessment of early ginglymodians

doi:10.19615/j.cnki.1000-3118.190319

Abstract

Abstract:

Ginglymodi are a subgroup of holostean fishes, including living gars and their closely-related fossil taxa. The early Middle Triassic (Anisian, ~244 Ma) kyphosichthyiforms from Yunnan and Guizhou, China represent the earliest records of this clade. Here, we report the discovery of a new kyphosichthyiform fish, Yudaiichthys eximius gen. et sp. nov., on the basis of four well-preserved specimens from the second (upper) member of Guanling Formation in Luoping, eastern Yunnan. The new discovery stimulated a phylogenetic analysis to reassess the interrelationships of the Kyphosichthyiformes and their relationships with other early ginglymodians. Results of our analysis indicate that the previously defined family Kyphosichthyidae and the genus Sangiorgioichthys are paraphyletic. A revised Kyphosichthyiformes is proposed here, and it is divided into two families, Kyphosichthyidae and Lashanichthyidae fam. nov. The family Kyphosichthyidae is restricted to include two genera Kyphosichthys and Fuyuanichthys. The Chinese “Sangiorgioichthys” species are removed into a new genus Lashanichthys, which is recovered as a taxon sister to Yudaiichthys gen. nov., and both genera are grouped into the new family, Lashanichthyidae. Sangiorgioichthys is restricted to include two species (S. aldae and S. valmarensis) from the late Middle Triassic (Ladinian) of the Monte San Giorgio area. The genus is removed out of the Kyphosichthyiformes and is recovered as the sister taxon of the Semionotiformes-Lepisosteiformes clade. The revised topology provides new insights into the anatomical evolution during the earliest ginglymodian history.

Key words: Luoping,Yunnan, Triassic, Kyphosichthyiformes, Ginglymodi, osteology, phylogeny

摘要：

铰齿鱼类是全骨鱼类中的一支,包括现生的雀鳝及其关系密切的化石类型。产自云贵地区中三叠世安尼期(~244 Ma)地层的拱鱼目(Kyphosichthyiformes)鱼类代表了铰齿鱼类最早的化石记录。根据最近在云南罗平关岭组二段发现的4块保存完好的鱼化石,命名了拱鱼目一个新属种,优美玉带鱼(Yudaiichthys eximius gen. et sp. nov.)。新的发现为重新研究拱鱼目和其他早期铰齿鱼类的系统发育关系提供了契机。分支分析结果表明,过去定义的拱鱼科(Kyphosichthyidae)和圣乔治鱼属(Sangiorgioichthys)都是并系类群。重新厘定后的拱鱼目分为拱鱼科和腊山鱼科(Lashanichthyidae fam. nov.); 其中,拱鱼科包括拱鱼属(Kyphosichthys)和富源鱼属(Fuyuanichthys); 苏氏圣乔治鱼(Sangiorgioichthys sui)和羊圈圣乔治鱼(S. yangjuanensis)归入新建的腊山鱼属(Lashanichthys gen. nov.), 和玉带鱼属一起组成腊山鱼科。圣乔治鱼属被移出拱鱼目,限定于产自圣乔治山地区中三叠世拉丁期地层的两个种(Sangiorgioichthys aldae和S. valmarensis); 该属和更进步的铰齿鱼类(半椎鱼目和雀鳝目)构成姐妹群关系。修订后的分支图为了解铰齿鱼类解剖特征的早期演化历史提供了新的见解。

关键词: 云南罗平, 三叠纪, 拱鱼目, 铰齿鱼类, 骨骼学, 系统发育

CLC Number:

Q915.862

XU Guang-Hui, MA Xin-Ying, WU Fei-Xiang, REN Yi. A Middle Triassic kyphosichthyiform from Yunnan, China, and phylogenetic reassessment of early ginglymodians. Vertebrata Palasiatica, 2019, 57(3): 181-204.

徐光辉, 马昕莹, 吴飞翔, 任艺. 2019, 57(3): 181-204, 云南中三叠世拱鱼目一新属种及早期铰齿鱼类系统发育关系再评估. 古脊椎动物学报.

Figures/Tables 8

References 72

[1]	Alvarado-Ortega J, Espinosa-Arrubarrena L, 2008. A new genus of ionoscopiform fish (Halecomorphi) from the Lower Cretaceous (Albian) lithographic limestones of the Tlayúa quarry, Puebla, Mexico. J Paleont, 82:163-175 DOI URL
[2]	Arratia G, 2013. Morphology, taxonomy, and phylogeny of Triassic pholidophorid fishes (Actinopterygii, Teleostei). Soc Vert Paleont Mem (Suppl J Vert Paleont), 13: 1-138
[3]	Bartram A W H, 1977. The Macrosemiidae, a Mesozoic family of holostean fishes. Bull Br Mus Geol, 29:137-234
[4]	Benton M J, Zhang Q Y, Hu S X et al., 2013. Exceptional vertebrate biotas from the Triassic of China, and the expansion of marine ecosystems after the Permo-Triassic mass extinction. Earth Sci Rev, 125:199-243 DOI URL
[5]	Brito P M, Alvarado-Ortega J, Meunier F J, 2017. Earliest known lepisosteoid extends the range of anatomically modern gars to the Late Jurassic. Sci Rep, 7:17830 DOI URL
[6]	Bürgin T, 2004. †Eosemionotus ceresiensis sp. nov., a new semionotiform fish (Actinopetygii, Halecostomi) from the Middle Triassic of Monte San Giorgio (southern Switzerland). In: Arratia G, Tintori A eds. Mesozoic Fishes 3 - Systematics, Pleoenvironments and Biodiversity. Munich: Verlag Dr. Friedrich Pfeil. 239-251
[7]	Cavin L, 2010. Diversity of Mesozoic semionotiform fishes and the origin of gars (Lepisosteidae). Naturwissenschaften, 97:1035-1040 DOI URL
[8]	Cavin L, Suteethorn V, 2006. A new Semionotiform (Actinopterygii, Neopterygii) from Upper Jurassic-Lower Cretaceous deposits of North-East Thailand, with comments on the relationships of Semionotiforms. Palaeontology, 49:339-353 DOI URL
[9]	Cavin L, Deesri U, Suteethorn V, 2013. Osteology and relationships of Thaiichthys nov. gen.: a Ginglymodi from the Late Jurassic-Early Cretaceous of Thailand. Palaeontology, 56:183-208 DOI URL
[10]	Cavin L, Deesri U, Veran M et al., 2018. A new Lepisosteiformes (Actinopterygii: Ginglymodi) from the Early Cretaceous of Laos and Thailand, SE Asia. J Syst Palaeont, doi: 10.1080/14772019.2018.1426060
[11]	Chen W Q, Sun Z Y, Tintori A et al., 2014. A new species of Sangiorgioichthys Tintori & Lombardo, 2007 (Actinopterygii; Semionotiformes) from the Pelsonian (Anisian, Middle Triassic) of Guizhou Province, South China. Neues Jahrb Geol Paläont, Abh, 273:65-74
[12]	Cope E D, 1872. Observations on the systematic relations of the fishes. Proc Am Assoc Adv Sci, 20:317-343
[13]	Deesri U, Lauprasert K, Suteethorn V et al., 2014. A new species of the ginglymodian fish Isanichthys from the Late Jurassic Phu Kradung Formation, northeastern Thailand. Acta Palaeont Pol, 59:313-331
[14]	Deesri U, Jintasakul P, Cavin L, 2016. A new Ginglymodi (Actinopterygii, Holostei) from the Late Jurassic-Early Cretaceous of Thailand, with comments on the early diversification of Lepisosteiformes in Southeast Asia. J Vert Paleont, 36:e1225747 DOI URL
[15]	Drucker E G, Lauder G V, 2002. Wake dynamics and locomotor function in fishes: interpreting evolutionary patterns in pectoral fin design. Integr Comp Biol, 42:997-1008 DOI PMID
[16]	Ebert M, 2018. Cerinichthys koelblae, gen. et sp. nov., from the Upper Jurassic of Cerin, France, and its phylogenetic setting, leading to a reassessment of the phylogenetic relationships of Halecomorphi (Actinopterygii). J Vert Paleont, 38:e1420071 DOI URL
[17]	Gibson S Z, 2013a. A new hump-backed ginglymodian fish (Neopterygii, Semionotiformes) from the Upper Triassic Chinle Formation of southeastern Utah. J Vert Paleont, 33:1037-1050 DOI URL
[18]	Gibson S Z, 2013b. Biodiversity and evolutionary history of †Lophionotus (Neopterygii: †Semionotiformes) from the western United States. Copeia, 2013:582-603 DOI URL
[19]	Grande L, 2010. An empirical synthetic pattern study of gars (Lepisosteiformes) and closely related species, based mostly on skeletal anatomy. The resurrection of Holostei. Copeia, 10(Suppl):1-871
[20]	Grande L, Bemis W E, 1998. A comprehensive phylogenetic study of amiid fishes (Amiidae) based on comparative skeletal anatomy: an empirical search for interconnected patterns of natural history. Soc Vert Paleont Mem (Suppl J Vert Paleont), 4: 1-690
[21]	Hu S X, Zhang Q Y, Chen Z Q et al., 2011. The Luoping biota: exceptional preservation, and new evidence on the Triassic recovery from end-Permian mass extinction. Proc R Soc B, 278:2274-2282 DOI URL
[22]	Lin H Q, Sun Z Y, Tintori A et al., 2011. A new species of Habroichthys Brough, 1939 (Actinopterygii; Peltopleuriformes) from the Pelsonian (Anisian, Middle Triassic) of Yunnan Province, South China. Neues Jahrb Geol Paläont, Abh, 262:79-89
[23]	Lombardo C, Tintori A, 2008. A new semionotid fish (Actinopterygii) from the Upper Triassic of northern Italy. In: Arratia G, Schultze H P, Wilson M V H eds. Mesozoic Fishes - Homology and Phylogeny. Munich: Verlag Dr. Friedrich Pfeil. 129-142
[24]	Lombardo C, Sun Z Y, Tintori A et al., 2011. A new species of the genus Perleidus (Actinopterygii: Perleidiformes) from the Middle Triassic of southern China. Boll Soc Paleont Ital, 50:75-83
[25]	Lombardo C, Tintori A, Tona D, 2012. A new species of Sangiorgioichthys (Actinopterygii, Semionotiformes) from the Kalkschieferzone of Monte San Giorgio (Middle Triassic; Meride, Canton Ticino, Switzerland). Boll Soc Paleont Ital, 51:203-212
[26]	López-Arbarello A, 2012. Phylogenetic interrelationships of ginglymodian fishes (Actinopterygii: Neopterygii). PLoS ONE, 7:e39370 DOI URL
[27]	López-Arbarello A, Alvarado-Ortega J, 2011. New semionotiform (Neopterygii) from the Tlayúa Quarry (Early Cretaceous, Albian), Mexico. Zootaxa, 2749:1-24 DOI URL
[28]	López-Arbarello A, Sferco E, 2018. Neopterygian phylogeny: the merger assay. R Soc Open Sci, 5:172337 DOI URL
[29]	López-Arbarello A, Wencker L C M, 2016. New callipurbeckiid genus (Ginglymodi: Semionotiformes) from the Tithonian (Late Jurassic) of Canjuers, France. Paläont Z, 90:543-560 DOI URL
[30]	López-Arbarello A, Sun Z Y, Sferco E et al., 2011. New species of Sangiorgioichthys Tintori and Lombardo, 2007 (Neopterygii, Semionotiformes) from the Anisian of Luoping (Yunnan Province, South China). Zootaxa, 2749:25-39 DOI URL
[31]	López-Arbarello A, Burgin T, Furrer H et al., 2016. New holostean fishes (Actinopterygii: Neopterygii) from the Middle Triassic of the Monte San Giorgio (Canton Ticino, Switzerland). PeerJ, 4:e2234 DOI URL
[32]	Ma X Y, Xu G H, 2017. A new ionoscopiform fish (Holostei: Halecomorphi) from the Middle Triassic (Anisian) of Yunnan, China. Vert PalAsiat, 55:92-106
[33]	Maisey J G, 1991. Araripelepidotes Silva Santos, 1985. In: Maisey J G ed. Santana Fossils: an Illustrated Atlas. Neptune, NJ: TFH Publications Inc. 118-123
[34]	Müller J, 1844. Über den Bau und die Grenzen der Ganoiden, und über das natürliche System der Fische. Ber Akad Wiss Berlin, 1844:416-422
[35]	Near T J, Eytan R I, Dornburg A et al., 2012. Resolution of ray-finned fish phylogeny and timing of diversification. Proc Nat Acad Sci, 109:13698-13703 DOI URL
[36]	Olsen P E, McCune A R, 1991. Morphology of the Semionotus elegans species group from the Early Jurassic part of the Newark Supergroup of eastern North America with comments on the family Semionotidae (Neopterygii). J Vert Paleont, 11:269-292 DOI URL
[37]	Patterson C, 1973. Interrelationships of holosteans. In: Greenwood P H, Miles R S, Patterson C eds. Interrelationships of Fishes. Zool J Linn Soc, 53(Suppl):233-305
[38]	Patterson C, 1982. Morphology and interrelationships of primitive actinopterygian fishes. Am Zool, 22:241-259 DOI URL
[39]	Regan C T, 1923. The skeleton of Lepidosteus, with remarks on the origin and evolution of the lower neopterygian fishes. Proc Zool Soc London, 1923: 445-461
[40]	Ruehl C B, Shervette V, Dewitt T J, 2011. Replicated shape variation between simple and complex habitats in two estuarine fishes. Biol J Linn Soc, 103:147-158 DOI URL
[41]	Schaeffer B, Dunkle D, 1950. A semionotid fish from the Chinle Formation, with considerations of its relationships. Am Mus Novit, 1457:1-29
[42]	Schröder K M, López-Arbarello A, Ebert M, 2012. Macrosemimimus gen. nov. (Actinopterygii, Semionotiformes) from the Late Jurassic of Germany, England and France. J Vert Paleont, 32:512-529 DOI URL
[43]	Schultze H P, Möller H, 1986. Wirbeltierreste aus dem Mittleren Muschelkalk (Trias) von Göttingen, West-Deutschland. Paläont Z, 60:109-129 DOI URL
[44]	Sun Z Y, Ni P G, 2018. Revision of Kyphosichthys grandei Xu & Wu, 2012 from the Middle Triassic of Yunnan Province, South China: implications for phylogenetic interrelationships of ginglymodian fishes. J Syst Palaeont, 16:67-85 DOI URL
[45]	Sun Z Y, Tintori A, Jiang D Y et al., 2009. A new perleidiform (Actinopterygii, Osteichthyes) from the middle Anisian (Middle Triassic) of Yunnan, South China. Acta Geol Sin, 83:460-470 DOI URL
[46]	Sun Z Y, Lombardo C, Tintori A et al., 2012. Fuyuanperleidus dengi Geng et al. 2012 (Osteichthyes, Actinopterygii) from the Middle Triassic of Yunnan Province, South China. Riv Ital Paleont Stratigr, 118:359-373
[47]	Sun Z Y, Lombardo C, Tintori A et al., 2015. A new species of Altisolepis (Peltopleuriformes, Actinopterygii) from the Middle Triassic of southern China. J Vert Paleont, 35:e909819 DOI URL
[48]	Sun Z Y, Tintori A, Lombardo C et al., 2016. New miniature neopterygians from the Middle Triassic of Yunnan Province, South China. Neues Jahrb Geol Paläont, Abh, 282:135-156
[49]	Swofford D L, 2003. PAUP. Phylogenetic analysis using parsimony (and other methods). Version 4.0b10. Sunderland, Massachusetts: Sinauer Associates
[50]	Tan K, Jin F, 2013. Re-study on Gymnoichthys inopinatus from Middle Triassic of Luoping, Yunnan, China. Vert PalAsiat, 51:1-16
[51]	Tintori A, Lombardo C, 2007. A new early Semionotidae (Semionotiformes, Actinopterygii) from the upper Ladinian of Monte San Giorgio area (southern Switzerland and northern Italy). Riv Ital Paleont Stratigr, 113:369-381
[52]	Tintori A, Sun Z Y, Lombardo C et al., 2007. New specialized basal neopterygians (Actinopterygii) from Triassic of the Tethys Realm. Geol Insubr, 10:13-20
[53]	Tintori A, Sun Z Y, Lombardo C et al., 2010. A new basal neopterygian from the Middle Triassic of Luoping County (South China). Riv Ital Paleont Stratigr, 116:161-172
[54]	Wen W, Zhang Q Y, Hu S X et al., 2012. A new genus of basal actinopterygian fish from the Anisian (Middle Triassic) of Luoping, Yunnan Province, Southwest China. Acta Palaeont Pol, 57:149-160
[55]	Wen W, Zhang Q Y, Hu S X et al., 2013. Coelacanths from the Middle Triassic Luoping Biota, Yunnan, South China, with the earliest evidence of ovoviviparity. Acta Palaeont Pol, 58:175-193
[56]	Wen W, Hu S X, Zhang Q Y et al., 2018. A new species of Platysiagum from the Luoping Biota (Anisian, Middle Triassic, Yunnan, South China) reveals the relationship between Platysiagidae and Neopterygii. Geol Mag, doi: 10.1017/S0016756818000079
[57]	Wenz S, 1967. Compléments à l’étude des poisons Actinoptérygiens du Jurassique français. Paris: Centre National de la Recherché Scientifique. 1-276
[58]	Wenz S, 1999. †Pliodetes nigeriensis, gen. nov. et. sp. nov., a new semionotid fish from the Lower Cretaceous of Gadoufaoua (Niger Republic): phylogenetic comments. In: Arratia G, Schultze H P eds. Mesozoic Fishes 2 - Systematics and Fossil Record. München: Verlag Dr. Friederich Pfeil. 107-120
[59]	Westoll T S, 1944. The Haplolepidae, a new family of Late Carboniferous bony fishes - a study in taxonomy and evolution. Bull Am Mus Nat Hist, 83:1-121
[60]	Wu F X, Sun Y L, Hao W C et al., 2009. New species of Saurichthys (Actinopterygii: Saurichthydae) from Middle Triassic (Anisian) of Yunnan Province, China. Acta Geol Sin, 83:440-450 DOI URL
[61]	Wu F X, Sun Y L, Xu G H et al., 2011. New saurichthyid actinopterygian fishes from the Anisian (Middle Triassic) of southwestern China. Acta Palaeont Pol, 56:581-614
[62]	Wu F X, Sun Y L, Fang G Y, 2018. A new species of Saurichthys from the Middle Triassic (Anisian) of southwestern China. Vert Palasiat, 56:273-294
[63]	Xu G H, Ma X Y, 2016. A Middle Triassic stem-neopterygian fish from China sheds new light on the peltopleuriform phylogeny and internal fertilization. Sci Bull, 61:1766-1774 DOI URL
[64]	Xu G H, Ma X Y, 2018. Redescription and phylogenetic reassessment of Asialepidotus shingyiensis (Holostei: Halecomorphi) from the Middle Triassic (Ladinian) of China. Zool J Lin Soc, 184:95-114 DOI URL
[65]	Xu G H, Shen C C, 2015. Panxianichthys imparilis gen. et sp. nov., a new ionoscopiform (Halecomorphi) from the Middle Triassic of Guizhou, China. Vert PalAsiat, 53:1-15
[66]	Xu G H, Wu F X, 2012. A deep-bodied ginglymodian fish from the Middle Triassic of eastern Yunnan Province, China, and the phylogeny of lower neopterygians. Chin Sci Bull, 57:111-118 DOI URL
[67]	Xu G H, Zhao L J, 2016. A Middle Triassic stem-neopterygian fish from China shows remarkable secondary sexual characteristics. Sci Bull, 61:338-344 DOI URL
[68]	Xu G H, Shen C C, Zhao L J, 2014a. Pteronisculus nielseni sp. nov., a new stem-actinopteran fish from the Middle Triassic of Luoping, Yunnan Province, China. Vert Palasiat, 52:364-380
[69]	Xu G H, Zhao L J, Coates M I, 2014b. The oldest ionoscopiform from China sheds new light on the early evolution of halecomorph fishes. Biol Lett, 10:20140204 DOI URL
[70]	Xu G H, Ma X Y, Ren Y, 2018a. Fuyuanichthys wangi gen. et sp. nov. from the Middle Triassic (Ladinian) of China highlights the early diversification of ginglymodian fishes. PeerJ, 6:e6054 DOI URL
[71]	Xu G H, Ma X Y, Zhao L J, 2018b. A large peltopleurid fish from the Middle Triassic (Ladinian) of Yunnan and Guizhou, China. Vert PalAsiat, 56:106-120
[72]	Zhang Q Y, Zhou C Y, Lü T et al., 2009. A conodont-based Middle Triassic age assignment for the Luoping Biota of Yunnan, China. Sci China Ser D-Earth Sci, 52:1673-1678 DOI URL