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Mid-Cretaceous Rudists (Bivalvia: Hippuritida) From [Papers in Palaeontology, 2015, pp. 1–24] MID-CRETACEOUS RUDISTS (BIVALVIA: HIPPURITIDA) FROM THE LANGSHAN FORMATION, LHASA BLOCK, TIBET by XIN RAO1,2,PETERW.SKELTON3,JINGENGSHA1,HUAWEICAI1 and YASUHIRO IBA4 1State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 39 East Beijing Road, Nanjing, 210008, China; e-mails: [email protected], [email protected], [email protected] 2University of Chinese Academy of Sciences, Beijing, 100039, China 3Department of Earth, Environment and Ecosystems, The Open University, Milton Keynes, MK7 6AA, UK; e-mail: [email protected] 4Department of Natural History Sciences, Hokkaido University, N10W8, Kitaku, Sapporo, Hokkaido 060-0810, Japan; e-mail: [email protected] Typescript received 9 March 2015; accepted in revised form 11 May 2015 Abstract: The Cretaceous Langshan Formation, which appearance. Associated Eoradiolites cf. davidsoni is of smal- crops out along the northern portion of the Lhasa block, is ler size and, by contrast, shows the celluloprismatic struc- composed mainly of grey or grey-black fine-grained lime- ture of the outer shell layer. The recently described stones, argillaceous limestones and a few thin-bedded canaliculate polyconitid genus, Magallanesia Sano et al., siltstones and silty mudstones. From specimens collected 2014, is represented by M. rutogensis sp. nov., which differs and/or photographed in the field, combined with a revision from the type species in possessing more numerous and of taxa previously recorded from the Langshan Formation, ventrally extended canals in the left valve. Possibly belong- four rudist taxa are recognized: (1) Auroradiolites gen. nov.; ing with the latter species are isolated right valves, similar (2) Eoradiolites cf. davidsoni; (3) Magallanesia rutogensis sp. to Horiopleura, but containing tabulae. A probable Sellaea nov.; and (4) Sellaea sp. We propose Auroradiolites gen. sp. is represented by poorly preserved right valves neverthe- nov. for the grouping of SW Asian to Pacific radiolitid spe- less showing typical features, including a very thin outer cies formerly attributed to Eoradiolites and characterized by shell layer and an erect posterior myophoral wall, separating a compact (non-celluloprismatic) outer shell layer. The new off a broad posterior ectomyophoral cavity. Based on the genus is represented in the Langshan Formation by presence of Magallanesia and the orbitolinid record, this A. biconvexus (Yang et al., 1982), including several other rudist fauna is probably no older than late Albian, while a synonymized taxa. Taxonomic confusion in the literature younger age cannot be excluded. Both Magallanesia and has resulted from the varied appearance of the strongly Auroradiolites are characteristic of a SW Asian/central developed ligamentary infolding in such forms: in the right Pacific faunal province, while the other two genera are valve, it has a forked inner termination, but common trans- more widely distributed. verse sections through both valves show the latter juxta- posed against its broad, rounded counterpart in the left Key words: Radiolitidae, Auroradiolites gen. nov., Polyconi- valve, giving the combined terminations a bulb-shaped tidae, orbitolinids, Albian, SW Asian – Pacific province. T HE elaborate morphologies of rudist bivalves (Cestari One such area of rudist colonization was the Lhasa and Sartorio 1995; Skelton 2013a) enabled them to block, which was located on the northern margin of the occupy shallow marine habitats in great abundance on eastern Tethys Ocean in mid-Cretaceous times (Chang the vast carbonate platforms that developed within the et al. 1986; Zhang 2000), and is today bounded on its equatorially encircling Tethyan–Atlantic–tropical Pacific northern side by the Bangong–Nu Suture and on its oceanic realm of the Late Jurassic and Cretaceous. There, southern side by the Indus–Yarlung Suture (XZBGM they congregated in loose to dense clusters, often forming 1993; Fig. 1). Massive mid-Cretaceous carbonate deposits broad shelly ‘meadows’ (Gili et al. 1995) or low mounds are exposed in the northern portion of the Lhasa block, sometimes preserved as laterally extensive biostromes to referred to as the Langshan Formation (Leier et al. 2007). bioherms within carbonate platform successions. This formation is distributed in a belt encompassing © The Palaeontological Association doi: 10.1002/spp2.1019 1 2 PAPERS IN PALAEONTOLOGY Rutog, Gar, Gegyai, Zhongba, Gerze, Coqen, Nyima, later to Eoradiolites and Sphaerulites (Scott et al. 2010), Xainza and Baingoin counties (XZBGM 1993; Fig. 1), and though possessing an entirely compact outer shell layer. it contains abundant orbitolinids, gastropods, corals and Similar forms were also reported from Iran, central rudists (Zhang 1982, 1986, 1991; Yu and Xia 1985; Loser€ Afghanistan, northern Pakistan, northern India and Japan and Liao 2001; Scott et al. 2010). (Montenat et al. 1982; Pudsey et al. 1985; Mathur and The rudists from the Langshan Formation have been Vogel 1988; Masse and Gallo Maresca 1997; Sano and described in Rutog, Gegyai, Gerze and Baingoin counties. Masse 2013; Upadhyay 2014 and see also Upadhyay 2001, Nineteen species have been described in the previous lit- fig. 4b). Yang et al. (1982) and Gou and Shi (1998) estab- erature, assigned to 11 genera (Yang et al. 1982; Yang lished many new species for these radiolitids based mainly 1984; Gou 1994; Gou and Shi 1998; Scott et al. 2010). on supposed differences in the form of their ligamentary However, in most cases, the diagnostic characters of the infolding in transverse section. But both Masse and Gallo genera or even families have not been well established in Maresca (1997) and Scott et al. (2010) questioned the these publications, so their identification remains prob- reliability of such variation in ligament form as a diag- lematic and in need of revision according to recent nostic criterion. Moreover, these south-west Asian and advances in rudist classification. Such revision is of par- western Pacific radiolitids differ from typical Eoradiolites ticular interest given the intermediate palaeogeographical (Pons et al. 2010) not only in the structure of the outer position of this fauna between the classically documented shell layer, but also in the shape of the shell and the myo- rudist faunas of the Mediterranean/Middle Eastern Teth- cardinal apparatus. yan region and those of Pacific provenance, largely Accordingly, these Lhasa block radiolitid species are described in the last few years (Masse and Shiba 2010; revised herein based on the investigation of material Sano and Masse 2013; Skelton et al. 2013; Sano et al. newly collected and/or photographed in the field by the 2014). senior author, and revision of the previous publications. The most abundant rudists from the Langshan Forma- In addition, three further rudist species are also described tion are some radiolitids that were originally assigned to for the first time from China and their stratigraphical and Praeradiolites (Yang et al. 1982; Gou and Shi 1998), but palaeogeographical significance clarified. FIG. 1. Map showing the outcrop of the Langshan Formation in the Lhasa block (modified from XZBGM, 1993). BNS, Bangong–Nu Suture; IYS, Indus–Yarlung Suture. RAO ET AL.: MID-CRETACEOUS RUDISTS FROM TIBET 3 GEOLOGICAL SETTING been deposited in the Gangdese retroarc foreland basin (Zhang et al. 2011). The rudist bivalves described herein were collected from The Langshan Formation crops out widely along a the limestone beds of the following three sections in belt encompassing Rutog, Gar, Gegyai, Zhongba, Gerze, Tibet: (1) the north side of Lameila Mountain at the Coqen, Nyima, Xainza and Baingoin counties, extending junction of Rutog County and Gar County (N from Rutog County in the west to Namtso Lake in the 32°42051″, E 79°51025″; Fig. 2A); (2) the south bank of east (Fig. 1; XZBGM 1993). It was deposited in a rela- Nyer Lake in Gegyai County (N 32°14031.5″,E tively stable carbonate sedimentary environment and is 82°07058.9″; Fig. 2B); and (3) the south-west bank of mainly composed of grey or grey-black fine-grained Zabuye Lake in Zhongba County (N 31°18016″,E limestones, argillaceous limestones and a few thin-bed- 83°55034″; Fig. 2C). ded siltstones and silty mudstones (Fig. 2D; Marcoux In the tectonic division of Tibet, these areas belong to et al. 1987; Zhang et al. 2004; Leier et al. 2007). The the northern part of the Lhasa block, which is sandwiched thickness of the Langshan Formation is poorly con- between the Bangong–Nu Suture and the Indus–Yarlung strained; we regard it as about 1000 m based on Zhang Suture (Fig. 1; Yin and Harrison 2000; Zhang 2000; Leier et al. (1986) and Yin et al. (1988). Based on its litholog- et al. 2007). Here, thick shallow marine carbonates of ical character, the Langshan Formation was separated mid-Cretaceous age, called the Langshan Formation, are into lower and upper parts, the former containing widely distributed. This formation is interpreted to have limestones, siltstones and marlstones, and the latter FIG. 2. The fossil localities in AC D Rutog (A), Gegyai (B) and Zhongba (C) counties, and the lithological features of the Langshan Formation and the beds yielding rudists (based on Zhang et al. 1986) (D). B 4 PAPERS IN PALAEONTOLOGY dominated by limestones (Zhang et al. 1986). The rudist Superfamily RADIOLITOIDEA d’Orbigny, 1847 specimens described herein were all collected from the Family RADIOLITIDAE d’Orbigny, 1847 upper part of the Langshan Formation (Fig. 2D). This formation conformably overlies the Lower Cretaceous si- Genus AURORADIOLITES gen. nov. liciclastic Duoba Formation, and is in turn unconform- ably overlain by the Upper Cretaceous coarse-grained LSID. urn:lsid:zoobank.org:act:AC0FD0E6-2E5C-4249-A935- clastic Jingzhushan Formation (BGMRXAR 1997; Zhao 77B57B7A0AA3 et al. 2001). Abundant marine fossils are present in the Langshan Type species. Praeradiolites gilgitensis Douville, 1926a; from the Formation and the main elements are orbitolinids, ru- upper Aptian of Kohistan, northern Pakistan. dists, gastropods and corals (Zhang 1982, 1986, 1991; Liao and Xia 1985; Yu and Xia 1985; BGMRXAR 1997; Derivation of name.
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