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Lingulata, Brachiopoda) from Southern China Zhifei Zhang,1 Jian Han,1 Xingliang Zhang,1 Jianni Liu1, Junfeng Guo1 and Degan Shu1,2

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Lingulata, Brachiopoda) from Southern China Zhifei Zhang,1 Jian Han,1 Xingliang Zhang,1 Jianni Liu1, Junfeng Guo1 and Degan Shu1,2 中国科技论文在线 http://www.paper.edu.cn Acta Zoologica (Stockholm) 88: 65–70 (January 2007) NoteBlackwell Publishing Ltd on the gut preserved in the Lower Cambrian Lingulellotreta (Lingulata, Brachiopoda) from southern China Zhifei Zhang,1 Jian Han,1 Xingliang Zhang,1 Jianni Liu1, Junfeng Guo1 and Degan Shu1,2 Abstract 1Department of Geology and Early Life Zhang Z., Han J., Zhang X., Liu J., Guo J. and Shu D. 2007. Note on the gut Institute, State Key Laboratory for preserved in the Lower Cambrian Lingulellotreta (Lingulata, Brachiopoda) Continental Dynamics, Northwest from southern China. — Acta Zoologica (Stockholm) 88: 65–70 University, Xi’an, 710069, China; 2School of Earth Sciences and Rescources, Lingulellotreta malongensis Rong is the earliest known taxon of the family China University of Geosciences, Beijing, Lingulellotretidae, which is characterized by the presence of a pedicle foramen 100083, China as well as an internal pedicle tube. New material from the Early Cambrian Chengjiang Lagerstätte of southern China provides improved anatomical Keywords knowledge for lingulellotretid species especially for the digestive system. linguloidea, Brachiopoda, digestive tract, Early Cambrian, Chengjiang Lagerstätte Additional gut fossils exhibit distinctly the anterior portion composed of esophagus and distended stomach, situated in the alleged visceral cavity, with Accepted for publication: the recurved intestine accommodated inside a hollow pseudointerarea. 2 August 2006 The frequency of occurrence of this intestinal layout suggests that this is not just an artefact of preservation. The gross configuration of the guts and the way they are preserved in the fossils suggest that they are in situ and, therefore, we can assume that Lingulellotreta had a hollow cavity presumably subtended by the pseuodelthyrium, which was invaded and occupied by the visceral organs. Hence, these fossils demonstrate the dangers of extrapolating crown-group soft-tissue configuration to the stem group. Z. F. Zhang, Early Life Institute and Department of Geology, Northwest University, Xi’an, 710069, China. E-mail: [email protected] or [email protected] Yunnan Province, southern China, which has surpassed the Introduction classic Burgess Shale for yielding important information on The Lingulata was originally established by Gorjansky several key steps in early deuterostome evolution (e.g. Shu and Popov (1985, 1986) as a separate class clade outside the et al. 2003, 2004, and references therein), as well as in the Brachiopoda. However, it is now commonly regarded as a preservation of nonmineralized internal tissues of known monophyletic group within the Brachiopoda (Holmer 1989; brachiopod taxa (Zhang et al. 2003, 2004a,b, 2005, 2006; Popov et al. 1993; Carlson 1995; Holmer et al. 1995; Cohen Holmer and Caron 2006). In this deposit, the family Lin- et al. 1998). The class then comprised three orders: Lingulida, gulellotretidae is represented by the numerically dominant Acrotretida and Siphontretida. In the revised Treatise on brachiopod Lingulellotreta malongensis Rong, 1974, which is Invertebrate Palaeontology (Part H) (Holmer and Popov the earliest known taxon of the family, and which is unique 2000), the concept of the Lingulida was expanded to encom- within the Linguloidea in having an elongate pedicle foramen pass the three superfamilies Linguloidea, Discinoidea and and an internal pedicle tube (Holmer et al. 1997). Details of Acrotheloidea. Of these, the first two are the only extant the morphology of the lingulellotretid species have been organophosphatic-shelled stocks, and thus have unquestion- made available (Jin et al. 1993; Holmer et al. 1997; Zhang able living representatives. They share many characters, et al. 2004a,b, 2005), including the pedicle, musculature, including type of musculature, pedicle, digestive tract, mantle mantle canal system, as well as the organization of the lopho- canal system, ontogeny, and shell structure. The majority of phore and the digestive system (Zhang et al. 2004a,b, 2005). these characters can also be traced back to their fossil repre- In this paper, we illustrate additional specimens from the sentatives (Steel-Petrovic 1976). Fossil lingulids are abundant Chengjiang Lagerstätte showing preservation of a U-shaped in the celebrated Early Cambrian Chengjiang Lagerstätte in digestive tract. The additional well-preserved material of © 2007 The Authors Journal compilation © 2007 The Royal Swedish Academy of Sciences 转载 中国科技论文在线 http://www.paper.edu.cn Note on the Lower Cambrian lingulid gut • Zhang et al. Acta Zoologica (Stockholm) 88: 65–70 (January 2007) Lingulellotreta allows a further account given here of the dictated by the activities of bacteria during early diagenesis, disposition of the alimentary canal inside the shells. which most likely varied in category of organism, colonizing either on or inside the tissue masses. However, this proposal remains to be resolved. Anyway, the three-dimensional ali- Preservation mentary canals suggest that the early diagenetic mineraliza- The strata containing the Chengjiang fossils come from the tion of some Chengjiang animals most likely took place prior Yu’anshan Member (Eoredlichia Zone), i.e. the upper part of to the breakdown of their nonmineralized carcasses. the Early Cambrian Heilinpu (formerly Qiongzhusi) Forma- tion, exposed in a wide area around Kunming, Yunnan, New fossil evidence China (Luo et al. 1999). The fossils in our collection are reddish-brown or yellowish-green after weathering, and The U-shaped arrangement of the digestive canal in Lingule- embedded in greyish-green and greyish-yellow mudstone llotreta malongensis has been confirmed by fossils from the intercalated with greyish-brown siltstone. The mudstone is Lower Cambrian of southern China (Zhang et al. 2004a). fine-grained enough for the details of the fossils to be pre- Based on analogy with extant species, the digestive tract served with remarkable fidelity. However, differential weath- should unquestionably be accommodated inside the visceral ering played an essential role in the final appearance of the cavity. Nevertheless, close examination of many more macroscopic Chengjiang fossils in that the fresh mudstone superbly well-preserved informative specimens (Fig. 1) from the strata is dark grey and that contrasts little with the urges us to cast doubt on this ubiquitous argument. As seen contained fossils, which make it difficult to differentiate a in Fig. 1(A), the alimentary canal is represented by a narrow, fossil from the surrounding matrix. However, a fossil, when partially mud-filled three-dimensional tube-like structure deeply weathered, appears as a variable degree of reddish- (Fig. 2A). It arises at the base of the less intricately spiral brown to yellowish-brown, with a striking colour contrast to lophophore, extends posteriorly as an approximately 0.5- to the surrounding greyish-green to greyish-yellow rock. 1-mm-long oesophagus leading into a pouch-like stomach The new specimens of Lingulellotreta examined here are all (Fig. 2A,B). Beyond the stomach, the digestive canal con- dorsoventrally compressed, and preserved parallel to the tinues as a narrow intestine filled with muddy sediment. bedding plane. Their shells are flattened into a thin film so Extending posteriorly approximately 0.8 mm away from the that the two valves are indistinguishable from each other. The stomach, the intestine markedly passes through the hinge line digestive tract of Lingulellotreta malongensis occurs either as a into the pseudointerarea, and continues as an approximately darkish thread-like line or partly or completely mud-filled 1.2-mm mud-filled tube (Fig. 2A,B). Then, it recurves and tubes with considerable relief (also see Zhang et al. 2004a). instead extends forwards close to the extremity of hinge line, However, the ingestion of sediment is an unlikely explana- finally terminating in an anal opening on the right anterior tion for the three-dimensional preservation of the alimentary body wall (Fig. 1A–C). canals in the Chengjiang lingulids. This is because brachio- In specimen ELI L-0060, composed of part and counter- pods are suspension filter-feeders. Thus, it would be difficult part, the digestive tract is a distinct darkish-brown recurved to explain the presence of a large amount of mud ingested in line (Fig. 1D,E). With respect of the disposition, this struc- their gut (Zhang et al. 2005). The preservation of soft tissues ture coincides well with the gut seen in Fig. 1(A–C), both indicates that these specimens represent complete carcasses invading and occupying the hollow cone-shaped interspace rather than shell valves. Although the exceptional preserva- presumably being subtended by the pseuodelthyrium, delin- tion of the Chengjiang fauna has attracted considerable eated by a triangular lineation in the fossils (Fig. 2). The attention, efforts have been focused on the various processes upward recurved portion of the intestine can be preserved in that triggered the preservation of the soft-bodied organisms many specimens (Fig. 1) (also see Zhang et al. 2004a, Fig. 1F). (Babcock et al. 2001; Zhu et al. 2001, 2005; Gabbott et al. Twenty fossils were available for the study presented here; all 2004). The underlying mechanisms for the fossilization of are deposited in the Early Life Institute (prefix ELI) and the soft tissues and hard skeletons remain obscure. Intriguingly, Department of Geology, Northwest University, Xian, China. both the slightly mineralized shells and the nonmineralized labile organic tissues
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