From the Lower Devonian in Qujing, Yunnan, China
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Historical Biology An International Journal of Paleobiology ISSN: (Print) (Online) Journal homepage: https://www.tandfonline.com/loi/ghbi20 Falxcornus, a new genus of Tridensaspidae (Galeaspida, stem-Gnathostomata) from the Lower Devonian in Qujing, Yunnan, China Xinyuan Meng & Zhikun Gai To cite this article: Xinyuan Meng & Zhikun Gai (2021): Falxcornus, a new genus of Tridensaspidae (Galeaspida, stem-Gnathostomata) from the Lower Devonian in Qujing, Yunnan, China, Historical Biology, DOI: 10.1080/08912963.2021.1952198 To link to this article: https://doi.org/10.1080/08912963.2021.1952198 Published online: 27 Jul 2021. Submit your article to this journal View related articles View Crossmark data Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=ghbi20 HISTORICAL BIOLOGY https://doi.org/10.1080/08912963.2021.1952198 Falxcornus, a new genus of Tridensaspidae (Galeaspida, stem-Gnathostomata) from the Lower Devonian in Qujing, Yunnan, China Xinyuan Menga,b and Zhikun Gaia,b,c aInstitute of Vertebrate Palaeontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China; bUniversity of the Chinese Academy of Sciences, Beijing, China; cCAS Center for Excellence in Life and Paleoenvironment, Beijing, China ABSTRACT ARTICLE HISTORY A new genus and species of galeaspid, Falxcornus liui gen. et sp. nov., is described from the Xishancun Received 25 April 2021 Formation (Lochkovian, Lower Devonian) in Qujing, Yunnan, China. The new taxon displays a suite of Accepted 27 June 2021 diagnostic characters of the Eugaleaspiformes and strikingly resembles the tridensaspids Tridensaspis and KEYWORDS Pterogonaspis in its sickle-like complex of cornual and inner cornual processes, leaf-like inner cornual Falxcornus; Lower Devonian; processes, and median dorsal opening posteriorly placed level with the centre of the orbital opening, but Galeaspida; Tridensaspidae; clearly differs from these genera in lacking the rostral process and laterally projecting cornual processes. An taxonomy; vertebrate extended phylogenetic analysis of the Galeaspida suggests that Falxcornus, Pterogonaspis and Tridensaspis palaeontology form a monophyletic group as the family Tridensaspidae, and Falxcornus is resolved as sister to Pterogonaspis and Tridensaspis. Therefore, Falxcornus liui gen. et sp. nov. now represents the most primitive and oldest known tridensaspid fish. Its subacute rostral margin and sickle-like complex of cornual and inner cornual processes likely represent an intermediate state of these highly specialised rostral and lateral projecting processes, respectively. The recurrent evolution of similar cephalic elaborations within the Galeaspida suggests that some galeaspids could manipulate water flow around their headshields to become active animals with higher manoeuvrability and versatility, similar to osteostracans. Introduction the Tridensaspidae by Shan et al. (2020), as it formed a monophyletic group together with Tridensaspis magnoculus in The Galeaspida isa clade of armoured jawless ‘ostracoderms’ (stem- all parsimony-based cladograms (Zhu 1992; Gai et al. 2005; Zhu gnathostomes) restricted to East Asia during the Middle and Gai 2006; Shan et al. 2020; Jiang et al. 2021). Ferrón et al. Palaeozoic (Tarlo 1967; Zhu 1992; Janvier 1996, 2009; Zhu and (2020,2021) showed that these cephalic elaborations of headshields Gai 2006; Gai et al. 2018; Shan et al. 2020; Jiang et al. 2021). They represent adaptations for passive hydrodynamic control both in the are among the enigmatic clades of stem-gnathostomes key to osteostracans and galeaspids. Therefore, the Tridensaspidae is an understanding the transition from jawless to jawed vertebrates important clade for understanding the last adaptive radiation of (Janvier 1996; Donoghue et al., 2000; Sansom et al., 2010; Gai galeaspids during the Pragian stage of the Early Devonian. et al., 2011; Gai and Zhu, 2012; Donoghue et al. 2014; Keating However, the evolution of cephalic elaborations within the and Donoghue 2016; Gai et al., 2019). Research on ostracoderm Eugaleaspiformes remains enigmatic because there is no intermedi morphology and phylogeny could facilitate interpretation of the ate state of these highly specialised rostral and lateral projecting gradual assembly of gnathostome characters such as jaws, processes. This condition is considerably different from that of the a mineralised dermal skeleton, vertebrae, a mineralised braincase, Huananaspiformes. Here, we describe a new eugaleaspidiform paired nostrils, a differentiated gut, a complex inner ear, externally genus and species from the lower part of the Xishancun open endolymphatic ducts, and paired pectoral and pelvic appen Formation (lower Lochkovian, Lower Devonian). The new taxon dages (Donoghue et al. 2014; Keating and Donoghue 2016; Gai et al. is characterised by a subacute rostral margin and sickle-like com 2011). Among galeaspids, the family Tridensaspidae plex of cornual and inner cornual processes, which likely represent (Eugaleaspiformes) is of particular interest because their head an intermediate condition between Tridensaspis and Nochelaspis. shields exhibit significant convergence with those of the As the new taxon was collected from the lower Lochkovian, lower Huananaspiformes in possessing similar cephalic elaborations than the other members of the Tridensaspidae, it sheds new light on (elongated rostra, lateral expansions or processes), which were the early evolution of the Tridensaspidae. previously regarded as diagnostic characters of the Huananaspiformes (Janvier 1975). The Tridensaspidae was erected by Liu (1986) based on Tridensaspis magnoculus from the bottom of Anatomical abbreviations the Yukiang Formation (Pragian, Lower Devonian) of Guangxi, as br.f, branchial fossa; br.o, branchial opening; c, cornual process; da, the rostral process and lateral projecting processes of this taxon dorsal aorta; dcm, dorsal commissure; ic, inner cornual process; ifc, were unknown in all other members of the Eugaleaspiformes. infraorbital canal; ldc, lateral dorsal canal; ltc, lateral transverse Pterogonaspis yuhaii (Zhu 1992) from the Xujiachong Formation canal;md.o, median dorsal opening;mdc, median dorsal canal; (Pragian, Lower Devonian) of Qujing, Yunnan was later referred to obr.c, oralobranchial chamber; oesc,oesophagus; orb, orbital CONTACT Zhikun Gai [email protected] Institute of Vertebrate Palaeontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China; University of the Chinese Academy of Sciences, Beijing, China; Institute of Vertebrate Palaeontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China; CAS Center for Excellence in Life and Paleoenvironment, Beijing, China; University of the Chinese Academy of Sciences, Beijing, China © 2021 Informa UK Limited, trading as Taylor & Francis Group Published online 27 Jul 2021 2 X. MENG AND Z. GAI opening; pi, pineal fossa; ro, rostral process; soc1, anterior supraor and Zhu 2010; Xue et al. 2018) (Figure 1(c)). The Xishancun bital canal; soc2, posterior supraorbital canal; pb.b, postbranchial Formation is mainly composed of hard and dense quartz sand bar; pb.w, postbranchial wall; vr, ventral rim. stone with yellowish green and black shale. A layer of greyish yellow fine-grained sandstone is in conformable or disconformable contact with the Silurian Yulongssu Formation at the bottom of Geological setting the Xishancun Formation, and the Xitun Formation conformably The material of Falxcornus liui gen. et sp. nov. was collected by overlies it (Lin et al. 1998; Ma et al. 2009). A large number of Professor Yuhai Liu in May 1980 under a water pipe on the south galeaspid, placoderm, sarcopterygian and actinopterygian fossils side of the road in front of the Qujing Maternal and Child Health have been found in the Xishancun Formation, and these constitute Care Hospital, Yunnan Province, China (Figure 1(a, b)). The an important early vertebrate fauna (Zhu et al. 2000; Zhao and Lower Devonian strata in eastern Yunnan Province are well devel Zhu 2010). Galeaspids described from the Xishancun Formation oped with a complete sequence, known as the Cuifengshan Group, include Eugaleaspis changi (Liu 1965, Liu 1980), Yunnanogaleaspis which is divided into four formations: the Xishancun, Xitun, major (Pan and Wang 1980), Nochelaspis maeandrine (Zhu 1992), Guijiatun and Xujiachong formations from bottom to top (Wang Polybranchiaspis liaojiaoshanensis (Liu 1965), P. minor, Laxaspis 1984; Zhu 1992; Liu 2002; Zhu and Gai 2006; Ma et al. 2009; Zhao qujingensis, L. yulongssus, Diandongaspis xishancunensis (Liu Figure 1. Maps of the fossil locality of Falxcornus liui gen. et sp. nov. (a, b) and the fossil fish-bearing lithological column (c) in Qujing, Yunnan Province, China. HISTORICAL BIOLOGY 3 1975), Pseudolaxaspis rostrata (Liu 1975; Gai et al. 2018), a subtriangular headshield with cornual and inner cornual processes, ‘Dongfangaspis qujingensis’ (Pan and Wang 1981), Damaspis var a longitudinal oval median dorsal opening, and a typical eugaleaspid- tus (Wang and Wang 1982), Nanpanaspis microculus (Liu 1965), type sensory canal system. It differs from the Shuyuidae, Altigibbaspis huiqingae (Liu et al. 2018), Siyingia perlatuspinosa (Si Sinogaleaspidae and Eugaleaspidae in its leaf-like inner cornual pro et al. 2015), Pentathyraspis pelta (Pan 1992) and Stephaspis dipter cess, differs from Shuyuidae and Sinogaleaspidae in its slit-like median iga (Gai and Zhu 2007). This