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1 Nanjing Institute of Geology and Paleontology, Academia Sinica, Nanjing 210008, China, <gxli{at}nigpas.ac.cn> and Department of Geological Sciences, Virginia Polytechnic Institute and State University, Blacksburg 24061, <xiao{at}vt.edu>
The Tannuolinidae includes only two known genera—Micrina Laurie, 1986 and Tannuolina Fonin and Smirnova, 1967, both from the Lower Cambrian. They are phosphatic bimembrate small shelly fossils, consisting of two types of sclerites—the mitral and sellate sclerites. These sclerites show basal-internal accretional growth, spaced growth lamellae, carinae (in mitral sclerites of Tannuolina) or teeth (in mitral sclerites of Micrina), and unevenly distributed pores and canals. The scleritome of tannuolinids is poorly known because they are often preserved as disarticulated sclerites. Here we describe Micrina xiaotanensis new species and Tannuolina zhangwentangi Qian and Bengtson, 1989 from the Lower Cambrian Shiyantou and Yu'anshan formations (upper Meishucunian to Qiongzhusian stages) at Xiaotan, eastern Yunnan, South China. The new material not only extends the geographic range of Micrina, previously known only in Australia, but also includes two composite specimens of T. zhangwentangi: one with a pair of dextral and sinistral mitrals juxtaposed along the apertural margin of their decrescent sides, and the other with a smaller sellate ontogenetically merged with the sella of a larger sellate. Perhaps as a result of being contacting or imbricating surfaces, the decrescent, sella, and duplicature sides are also characterized by negative allometry (relative to other sides) and a sparse distribution of pores. The new fossils do not support reconstructions that place a sellate and a mitral sclerite, respectively, at the anterior and posterior end of the Tannuolina animal, in a way similar to Halkieria evangelista Conway Morris and Peel, 1995. Instead, they are consistent with the traditional view that the scleritome of Tannuolina consisted of four anterior–posterior files of sclerites: two opposing mitral rows flanked by two imbricated sellate series. The bimembrate Micrina may or may not have had a similar scleritome. If Tannuolina and Micrina form a monophyletic group outside the total group of brachiopods, then Micrina is likely to have had a multisclerite scleritome similar to that of Tannuolina. This implies that microstructural similarities between the tannuolinids and some linguliformean brachiopods are probably symplesiomorphic or convergent. Alternatively, Tannuolina and Micrina may represent stem groups leading to the linguliformeans while Halkieria represents a stem group leading to the calcareous brachiopods—a radical hypothesis that would imply that the two brachiopod groups independently evolved the bivalved body plan.
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