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1400 Van Iten.Vp Conulariids and Sphenothallus (Cnidaria, Medusozoa) from the Tonggao Formation (Lower Ordovician, China) HEYO VAN ITEN, LUCY A. MUIR, JOSEPH P. BOTTING, ZHANG YUAN-DONG & JIH-PAI LIN Conulariella sp. and possibly one other conulariid species occur in close association with Sphenothallus sp. in the lower part of the early Floian Tonggao Formation near the town of Sandu, Guizhou Province, China. This is the only known occurrence of Conulariella in Early Ordovician rocks outside of Bohemia (Perunican terrane), and also the first report of Sphenothallus from the Ordovician of China. The apertural margin of Conulariella appears to have been mostly straight, and apertural lappets probably were not present in this genus. Some Tonggao conulariids may have attached to orbiculoid brachiopods in life. Based on the most recent palaeogeographical reconstructions of Gondwana and associ- ated terranes, Conulariella could have had a dispersal path along the Gondwanan margin. • Key words: Conulariella, Sphenothallus, conulariids, Ordovician, South China. VAN ITEN, H., MUIR, L.A., BOTTING, J.P., ZHANG, Y.D. & LIN, J.P. 2013. Conulariids and Sphenothallus (Cnidaria, Medusozoa) from the Tonggao Formation (Lower Ordovician, China). Bulletin of Geosciences 88(4), 713–722 (5 fig- ures). Czech Geological Survey, Prague. ISSN 1214-1119. Manuscript received December 5, 2012; accepted in revised form July 22, 2013; published online August 14, 2013; issued October 31, 2013. Heyo Van Iten (corresponding author), Department of Geology, Hanover College, Hanover, IN 47243, U.S.A; and Cincinnati Museum Center, Research Associate, Department of Invertebrate Paleontology, 1301 Western Avenue, Cincinnati, OH 45203, U.S.A.; [email protected] • Lucy A. Muir, Joseph P. Botting, Zhang Yuan-Dong & Jih-Pai Lin, State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Acad- emy of Sciences, 39 East Beijing Road, Nanjing 210008, China; [email protected], [email protected], [email protected], [email protected] Conulariids (Cnidaria, Scyphozoa; Van Iten et al. 2006) are 2004). More recently, Pillet & Beaulieu (1998) described found in the geological record from the Ediacaran to the C. purkynei, C. minima Pillet & Beaulieu and Conulariella Triassic (Leme et al. 2008), and the related Sphenothallus sp. from the Middle Ordovician (Darriwilian) Schistes à (Cnidaria, Scyphozoa or Hydrozoa; Van Iten et al. 1992) is nodules d’Angers of the Armorican Massif (northwestern known from the Cambrian to the Permian (Fatka et al. France). The small shelly fossil Hexaconularia He & Yang 2012). Conulariids generally are rare in the Ordovician, and 1986, currently known only from basal Cambrian strata of they appear to exhibit marked faunal provincialism (Van South China (Conway Morris & Chen 1992), appears to be Iten & Vyhlasová 2004). In this paper we describe Spheno- most similar to Conulariella (Van Iten et al. 2010), and in- thallus sp. and two conulariids, Conulariella sp. and an in- deed these two genera may be nearest relatives. determinate conulariid, from the Lower Ordovician Tong- The present study constitutes the first definite report gao Formation of Guizhou Province, South China. of Conulariella in China as well as the first occurrence of Conulariella Bouček, 1928, originally described from this conulariid genus in Early Ordovician strata outside of Early Ordovician strata of Bohemia, is a highly distinctive Bohemia (Perunica). This is also the first report of genus characterized by (1) a strongly rectangular trans- Sphenothallus in the Ordovician of China. As well as ex- verse cross section and (2) nearly rectilinear transverse ribs panding our knowledge of the stratigraphical and palaeo- lacking any definition of a facial midline (e.g., Moore & geographical distribution of these two medusozoan gen- Harrington 1956a, fig. 45). Bouček (1928) recognized era, the new material from China provides additional three species, namely the type species C. robusta evidence regarding the locations of conulariid-bearing (Barrande, 1867), C. purkynei (Želízko, 1911) and C. sulca terranes during Early Ordovician times and/or the dis- (Želízko, 1921). All three species range from the earliest persal capacity and life history of conulariids and Floian to the latest Darriwilian (Van Iten & Vyhlasová Sphenothallus. DOI 10.3140/bull.geosci.1400 713 Bulletin of Geosciences Vol. 88, 4, 2013 Figure 1. Locality map showing the Xiayangao section through the Tonggao Formation (Lower Ordovician, lower Floian). Inset map shows the loca- tion of the section within China. Material and methods Stratigraphical and palaeoenvironmental context The present study is based on direct examination of 51 co- nulariid and 18 Sphenothallus specimens from a single sec- The rock exposure from which the Tonggao Formation co- tion through the Tonggao Formation near the town of nulariids and Sphenothallus were obtained is located on the Sandu, Guizhou Province, China (Fig. 1; see also discus- track between Xiayangao and Shijiawan villages, near the sion below). The specimens occur in blue-gray shale or town of Sandu, Guizhou Province, China (GPS co- yellow-green shale or siltstone, and were subjected to mi- ordinates N 26° 01.814´E 107° 48.637´; Fig. 1). Approxi- nimal preparation and cleaning with water and a soft brush. mately 80 m of blue-gray shale and yellow-green shale and Specimens were examined and photographed using reflec- siltstone of the Tonggao Formation is exposed along the ted light and scanning electron microscopy. Backscattered track, of which the lowest 30 m contains exceptionally pre- and secondary SEM imaging was performed using a Ge- served fossils (Muir et al. in press) together with shelly mini Leo 1530VP Environmental Scanning Electron Mic- taxa including conulariids and Sphenothallus. The rocks roscope and a Hitachi S-3000N Scanning Electron Micro- are of early Floian age (Zhang in Zhan & Jin 2008), with scope. Operating voltage in backscattered electron mode the sampled beds occurring in the Tetragraptus approxi- was 15–20 kV. Preservation of conulariid and Sphenothal- matus Biozone (Fig. 1). lus shell material was studied using a combination of More than 50 species of invertebrates and algae have so EDAX and elemental mapping. EDAX analysis was done far been collected from this section in the Tonggao Forma- using an Inca X-sight device made by Oxford Instruments. tion (Muir et al. 2011). The most abundant organisms are Light photography was performed with a Nikon D80 digi- rhynchonelliformean and linguliformean brachiopods, dis- tal SLR camera, combined with a Sigma 105 mm f2.8 articulated echinoderms (cystoids and possible eocrinoids macro lens and extension tubes, utilising direct sunlight or crinoids), dendroid graptolites, graptoloids, trilobites where possible and external, reflected flash at higher mag- and bivalved arthropods. Gastropods, tergomyans, bryo- nification. All specimens have been deposited in the Nan- zoans, hyoliths and sponge spicules are also present, but in jing Institute of Geology and Palaeontology (NIGP), speci- low numbers. A variety of trace fossils (horizontal burrows men numbers NIGP 156022 to NIGP 156067. up to 5 mm wide) and agglutinated tubes occur at several 714 Heyo Van Iten et al. Conulariids and Sphenothallus from the Lower Ordovician of China levels. Less easily preservable taxa include algae, palae- Conulariella sp. oscolecidan worms and a possible nematode (Muir et al.in Figure 2A–F press), and non-biomineralizing arthropods. The taphonomy of the Tonggao fossils varies accord- Horizon and locality. – Tonggao Formation (Lower Ordo- ing to their original composition, and overall the vician, lower Floian, Tetragraptus approximatus Bio- taphonomic style is a mixture of organic, mouldic and zone); near the town of Sandu, Guizhou Province, People’s early diagenetic iron-mineral preservation. Originally Republic of China. phosphatic groups (conulariids, Sphenothallus, phos- phatic brachiopods and palaeoscolecids) are preserved as Material. – Forty-seven nearly complete and partial speci- flattened moulds. Elemental mapping of conulariids has mens (NIGP 156022 to NIGP 156041, NIGP156059 to shown that the original phosphate has been dissolved, and NIGP156067). in some specimens there is a thin coating of weathered iron minerals replacing it. Description. – Specimens flattened, with the major faces Zhan & Jin (2008) interpreted the Tonggao Formation nearly pressed against each other and the minor faces lar- as being deposited in a deep-water environment becoming gely obscured; nearly complete specimens (Fig. 2B) gently shallower towards the top of the formation, based on the curved, ranging from approximately 7 mm long and brachiopod faunas. The presence in the studied section of 2.5 mm wide (maximum single face width) to approxima- numerous small (millimetre-scale) clay mineral concre- tely 24 mm long and 12 mm wide (maximum single face tions in the beds above those sampled for fossils indicates width), bluntly terminated at the apical end (apex is mis- an environment that was restricted and hypersaline, al- sing) but with a minimum single face width as low as ap- though deep-water (Izawa et al. 2012). However, the fos- proximately 0.2 mm (Fig. 2E); some partial specimens ori- sil-bearing beds were deposited in a more standard marine ginally at least 50 mm long. Apertural margin nearly environment; the echinoderms and other groups indicate straight (Fig. 2A, B). Angle of expansion
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