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Paleoecology of a Pennsylvanian Encrusting Colonial Rugose Coral in South Guizhou, China

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Paleoecology of a Pennsylvanian Encrusting Colonial Rugose Coral in South Guizhou, China Palaeogeography, Palaeoclimatology, Palaeoecology 280 (2009) 507–516 Contents lists available at ScienceDirect Palaeogeography, Palaeoclimatology, Palaeoecology journal homepage: www.elsevier.com/locate/palaeo Paleoecology of a Pennsylvanian encrusting colonial rugose coral in south Guizhou, China Yong-li Zhang a, En-pu Gong a,⁎, Mark A. Wilson b, Chang-qing Guan a, Bao-liang Sun a, Hong-lun Chang a a Department of Geology, Northeastern University, Shenyang, 110004, PR China b Department of Geology, The College of Wooster, Wooster, OH 44691, USA article info abstract Article history: The well-preserved Pennsylvanian encrusting colonial rugose coral Ivanovia is widespread and easily Received 4 February 2009 observed in south Guizhou, China. There are three common types of hard substrate encrusted by Ivanovia: in Received in revised form 20 June 2009 situ carbonate hardgrounds, carbonate hardground clasts, and calcareous bioclasts. Thin, spreading sheets are Accepted 1 July 2009 the most common growth form of Ivanovia in the study region. Ivanovia employed peripheral, medial and Available online 30 July 2009 mixed growth strategies to occupy a sufficient living space on the substrate. It favored a shallow, warm, and clear marine environment within the photic zone and had a high tolerance of water movement. Ivanovia was Keywords: generally smothered and covered by mud in the studied area. The Ivanovia fossil communities in south Hard substrate Encrusting Guizhou are characterized by a low species diversity conforming to the typical evolutionary pattern of hard Colonial rugose coral substrate marine communities in the Carboniferous. Paleoecology © 2009 Elsevier B.V. All rights reserved. Pennsylvanian South Guizhou 1. Introduction Lescinsky, 1997; Nakazawa, 2001). More work is necessary to better delineate the diversity pattern of hard substrate communities in the Until recently, encrusting metazoans were reported as relatively geologic record, so we need detailed studies of encrusting assem- rare on hard substrates in the Carboniferous (Alvarez and Taylor, 1987; blages in the Carboniferous (Brett, 1988; Taylor and Wilson, 2003). Brett, 1988; Lescinsky, 1997; Sumrall et al., 2006; Wilson and Palmer, The present study focuses on well-preserved examples of the 1992). In the Paleozoic, much of the increased abundance and Pennsylvanian encrusting colonial rugose coral Ivanovia. Specimens of diversity of hard substrate communities was mainly due to the this coral are widespread and easily studied in south Guizhou, China increase in hard substrate availability (Palmer, 1982; Wilson et al., (Guan et al., 2006). Encrusters are generally preserved in situ, 1989; Wilson et al., 1992; Wilson and Palmer, 1992; Taylor and Wilson, retaining their original positions on the substrate and thus offering 2003). Carbonate hardgrounds were rare in the Carboniferous (Suchy important advantages for a paleoecological study. The purpose of this and West, 1988; Wilson and Palmer, 1992; Taylor and Wilson, 2003) paper is to study the habit, growth forms and ecology of Ivanovia.It and thus the hard substrate niche was relatively uncommon at this provides a new example for the study of encrusting organisms on hard time. It is possible that the environmental stresses provided a selective substrates of the Carboniferous. pressure favoring marine benthic organisms which did not require hard substrates. In general, the low-diversity Carboniferous hard 2. Setting and stratigraphy substrate communities may have been overlooked because few fi workers expect to nd them (Lescinsky, 1994; Lescinsky, 1997). South Guizhou, southern China, is characterized by extended Although carbonate hardgrounds seem to be rare in the Carboniferous, outcrops of Carboniferous and Permian carbonate rocks. There are some other types of hard substrate such as calcitic skeletons, shells abundant fossils in these rocks, thus the region has been a focus of and mobile rocks are abundant. Studies of Carboniferous inorganic Carboniferous stratigraphy and paleontology (Wu, 1987; Li et al.,1996; hard substrates are thus surprisingly few (Palmer, 1982; Edwards and Liu et al., 1994; Feng et al., 1999; Jin et al., 2000; Wang and Qi, 2002; Marcus, 1993; Webb, 1993), but the encrusting taxa on organic hard Zhang, 2002). The Houchang region, south Guizhou, lies on the substrates are relatively well known from this interval (Taylor, 1985; southern margin of the Yangtze Plate, with its strata belonging to the Cossey and Mundy, 1990; Powers and Ausich, 1990; Molineux, 1994; Yunnan–Guizhou–Guangxi subregion of south China and southern Guizhou, northern Guangxi and southeastern Yunnan areas (Wang et al., 1990). Strata of Carboniferous to Permian age in this region are ⁎ Corresponding author. Tel.: +86 24 83675066; fax: +86 24 83689266. E-mail addresses: [email protected] (Y. Zhang), [email protected] (E. Gong), mostly light grey, medium- to thick-bedded and massive limestones [email protected] (M.A. Wilson). representative of shallow epicontinental sea deposits (Wu, 1987; 0031-0182/$ – see front matter © 2009 Elsevier B.V. All rights reserved. doi:10.1016/j.palaeo.2009.07.004 508 Y. Zhang et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 280 (2009) 507–516 Fig. 1. (A) Location of the studied area west of Houchang, south Guizhou Province, China. (B) A geological map of the study area. Y. Zhang et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 280 (2009) 507–516 509 GGMRB, 1987). The Carboniferous sequences consist of the fossilifer- high and about 10 cm long, was found on a fusulinid grainstone ous Weining and Maping formations (GGMRB, 1987). In the substrate (Fig. 4E). Pennsylvanian, this region was on the high-energy margin of a carbonate platform with many reefal buildups (Fan and Rigby, 1994; 4. Types of hard substrate encrusted by Ivanovia Guan et al., 2004; Gong et al., 2004; Guan et al., 2006, 2007; Sun et al., 2007; Gong et al., 2007a,b,c; Zhang et al., 2007). 4.1. Carbonate hardgrounds (in situ) The colonial rugose coral Ivanovia is abundant near the village of Bianping, about 4 km west of Houchang, Ziyun County, in Guizhou Hardgrounds are synsedimentarily lithified seafloors formed by Province (Fig. 1). It is found in a thick, cyclic sequence of bedded and precipitation of carbonate cements in primary pore spaces (Wilson massive Pennsylvanian and Permian limestones which also contain and Palmer, 1992). The substrate of Ivanovia in this study consists of large numbers of other benthic organisms such as fusulinids, corals, bioclastic grainstone with common fragments of fusulinids, crinoids and brachiopods. Ivanovia occurs in the Triticites Zone of the Maping and brachiopods, and of boundstone. The primary pore spaces Formation (uppermost Pennsylvanian) (Fig. 2)(Guan et al., 2006). between skeletal grains are filled with synsedimentary marine The common fusulinids include: Triticites pygmaeus, T. simplex, T. cements (botryoidal, isopachous and fibrous) (Fig. 5A, B and C). parvus, T. minimus, T. chinensis, T. subrhomboides, Schwagerina vul- Cement crusts (multi-generations of syndepositional calcite cement) garis, S. subnathorsti, S. kuenchinensis, and S. amushanensis (Ding are mainly developed in the larger cavities (Fig. 5D). The synsedi- et al., 1992). Brachiopods (Choristites sp., Squamularia sp., Echinaria mentary cements represent over 20–30% of the rock volume of the sp., Wellerella sp., Martinia sp.) and corals (Caninia sp., Fomitchevella substrate colonized by Ivanovia. Generally Ivanovia grew by following sp., Antheria sp., Ivanovia sp.) are also common in this sequence (Li the shape of the substrate, demonstrating that it was initiated on the and Zhao, 1993; Zhang, 2002). hardground after cementation (Fig. 5E). Tubiphytes is relatively abundant in the substrate underlying Iva- novia. Tubiphytes is generally considered to have been a binding 3. Morphology and growth forms of Ivanovia organism, but it also seems to have functioned as framework constructor (Riding and Guo, 1992; Senowbari-Daryan and Flügel, Ivanovia is a massive aphroid coral (Fig. 3). The corallites are 1993; Wang et al., 1994; Wang et al., 1998; Tian and Fan, 2001; Shen connected with cystosa. Major septa are long, with their bases and Xu, 2005). In the study region, Ivanovia occasionally colonized disrupted by cystosepiments; they rarely reach the columella. Minor Tubiphytes boundstone. Tubiphytes is often associated with bioclasts septa are occasionally developed. The columella is a thin plate or cemented by synsedimentary cement in the substrate underneath fusuliform. Tabulae lightly incline toward the center. The internal Ivanovia (Fig. 6). Tubiphytes boundstone contains some tubular structural organization of Ivanovia is aphroid (Fig. 3A); the astogeny of structures filled with calcite lacking any internal features, suggesting this coral involves a form of non-parricidal budding (Scrutton, 1998). that they may be borings. Thin, spreading sheets are the most common growth form of Ivanovia In the study region, the outcrops with Ivanovia encrusting the in the study region (Fig. 4A). These colonies generally are 2–10 cm carbonate hardgrounds have a lateral extent of 1–10 m. The Ivanovia thick and 1–10 m long. Ivanovia also has both spreading tabular and beds are discontinuous so that none of the hardgrounds encrusted by domal growth forms with a maximum colony thickness of 30–40 cm Ivanovia forms continuous, unbroken platforms; they are all con- (Fig. 4B). Occasionally, thin, spreading sheets, which grew
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