Body-size changes of latest Permian brachiopods in varied palaeogeographic settings in South China and implications for controls on animal miniaturization in a highly stressed marine ecosystem
Introduction
Body size is a key, intrinsic character of any organism and profoundly affects its biology and ecology (Calder, 1984, Jablonski, 1996). The study of body-size changes in space and time is important for understanding macro-evolution of organisms and ecosystems; it also provides important insights into the evolution history and dynamics of past environments. The commonly proposed controls on body-size changes include oxygen fluctuations (Savrda and Bottjer, 1986, Payne et al., 2008, Payne et al., 2013), food availability (Hallam, 1965, Rheault and Rice, 1996, Twitchett, 2007, He et al., 2010) and temperature changes (Hunt et al., 2010, Sheridan and Bickford, 2011, Edeline et al., 2013), as well as substrate conditions. As many of these factors vary with water depth, the relationship between body size and bathymetry is a critical aspect of studies on body-size changes (Anderson, 1971, Thiel, 1975, Peck and Harper, 2010, Shi et al., 2016). Numerous researches have been undertaken on the Permian-Triassic body-size changes of conodonts, brachiopods, siliceous sponges, ostracods or foraminifers in South China (He et al., 2007a, He et al., 2010, He et al., 2015a, He et al., 2016, Peng et al., 2007, Luo et al., 2008, Song et al., 2011, Liu et al., 2013, Chu et al., 2016, Zhang et al., 2016). Most of these studies have focused on the patterns of body-size changes of individual taxa across time and their possible underlying control mechanisms, and a few (e.g., He et al., 2010, Liu et al., 2013) have addressed the relationship between size changes in relation to primary productivity and redox palaeoproxies in the context of palaeobathymetry. Most recently, based on a global dataset of Changhsingian brachiopod orders, Shi et al. (2016) examined the relationship of Changhsingian brachiopod body-size changes in relation to the onshore-offshore-basin gradient.
The present paper aims to investigate body-size changes of latest Permian brachiopods across different bathymetrically controlled palaeoenvironmental settings within the South China basin. For this purpose, we chose two most commonly found Changhsingian chonetid brachiopod species, Tethyochonetes pigmaea and Tethyochonetes quadrata, from six different sections, which together constituted an approximately-defined basinwide bathymetric gradient spanning the shallow-water clastic shelf, shallow-water carbonate platform and ramp, and deep-water siliceous basinal settings (see below).
Section snippets
Geographical location, palaeogeographical setting and classification on depth-related palaeoenvironments
The studied sections include Zhongzhai, Huangzhishan, Daoduishan, Majiashan, Rencunping and Dongpan sections in South China. Geographically, Huangzhishan is located 12 km west of Huzhou City, Zhejiang Province (Fig. 1A). Daoduishan is located about 2.5 km northeast of the Meishan Section D of Changxing County, Zhejiang Province, where the GSSP of the Permian-Triassic Boundary (PTB) has been defined (Yin et al., 2001) (Fig. 1A). Majiashan is located about 55 km southeast of Hefei City, Anhui
Age analysis and correlation
At Zhongzhai, the PTB has been placed between Beds 29 and 30 (Fig. 3), according to the U-Pb age of 252.24 ± 0.13 Ma in Bed 29, which is basically equivalent to the calibrated PTB age of 252.17 ± 0.06 Ma in Bed 27c at the GSSP section of Meishan (Shen et al., 2011, Zhang et al., 2014). Beds 4 to 28 of this section have been assigned to the Changhsingian, based on the abundance of brachiopod Tethyochonetes pigmaea (typical for the Changhsingian in South China, see Shen and Archbold, 2002, Zhang et
Methods of brachiopod collection, body size measurement and definition
The studied specimens for Tethyochonetes pigmaea and T. quadrata were mainly collected from the Changhsingian (or the upper Changhsingian) at the studied sections (Table 1). All related brachiopods found during excavating were collected, including complete and incomplete individuals, but only the complete individuals of Tethyochonetes pigmaea and T. quadrata were chosen for body-size measuring for the purpose of this paper.
The width of each brachiopod individual refers to the maximum shell
Body-size profiles of T. pigmaea and T. quadrata through studied sections
The mean sizes (X) for T. pigmaea from Beds 18 to 36 of Huangzhishan fluctuated significantly, from 2.88 mm to 7.08 mm, while that of T. quadrata from the same section first slightly fluctuated from 6.22 mm to 7.26 mm from Beds 18 to 24, and then became reduced to the end of Changhsingian (Table 1, Fig. 3).
The mean sizes (X) for T. pigmaea from Beds 4 to 27-6 at Zhongzhai slightly fluctuated, from 4.50 mm to 5.38 mm, while the size profile for T. quadrata from Beds 4 to 27-5 of the same section
Taphonomic features of brachiopods
Brachiopods collected from Huangzhishan and Zhongzhai lack any evidence of abrasion. They are randomly distributed on bedding surfaces without any preferred orientation; they are abundantly preserved with articulated valves, indicating very limited or no post-mortem transportation (Chen and Shi, 1999, Chen et al., 2010). Brachiopods collected from Daoduishan have both complete and incomplete valves, but these valves are randomly-arranged on bedding surface, suggesting limited transportation
Acknowledgments
We wish to express sincere thanks to two reviewers for offering good comments and suggestions for the revision of this manuscript. This paper has been supported by NSFC (Grant Nos. 41372030, 40872008, 40502001), the Foundation of the Geological Survey of China (1212011220529), the Ministry of Education of China (B08030 of 111 Project, NCET-10-0712), and an Australian Research Council (ARC DP150100690) research grant to GRS.
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