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Size Variation of Conodont Elements of the Hindeodus–Isarcicella Clade During the Permian–Triassic Transition in South China and Its Implication for Mass Extinction

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Size Variation of Conodont Elements of the Hindeodus–Isarcicella Clade During the Permian–Triassic Transition in South China and Its Implication for Mass Extinction Palaeogeography, Palaeoclimatology, Palaeoecology 264 (2008) 176–187 Contents lists available at ScienceDirect Palaeogeography, Palaeoclimatology, Palaeoecology journal homepage: www.elsevier.com/locate/palaeo Size variation of conodont elements of the Hindeodus–Isarcicella clade during the Permian–Triassic transition in South China and its implication for mass extinction Genming Luo a, Xulong Lai a,⁎, G.R. Shi b, Haishui Jiang a, Hongfu Yin a, Shucheng Xie c, Jinnan Tong c, Kexin Zhang a, Weihong He c, Paul B. Wignall d a Faculty of Earth Science, China University of Geosciences,Wuhan 430074, PR China b School of Life and Environmental Sciences, Deakin University, 221 Burwood Hwy, Burwood VIC 3125, Australia c Key Laboratory of Geobiology and Environmental Geology, China University of Geosciences, Wuhan 430074, PR China d School of Earth and Environment, University of Leeds, Leeds. LS2 9JT, United Kingdom ARTICLE INFO ABSTRACT Article history: Based on the analysis of thousands of conodont specimens from the Permian –Triassic (P–T) transition at Meishan Received 17 August 2007 (the GSSP of P–T Boundary) and Shangsi sections in South China, this study investigates the size variation of Received in revised form 1 April 2008 Hindeodus and Isarcicella P1 elements during the mass extinction interval. The results demonstrate that Hin- Accepted 3 April 2008 deodus–Isarcicella underwent 4 episodes of distinct size reduction during the P–T transition at the Meishan Section and 2 episodes of size reduction in the earliest Triassic at Shangsi. The size reductions at Meishan took Keywords: Multi-episode mass extinction place at the junctions of beds 24d/24e, 25/26, 27b/27c and 28/29, and at the junctions of beds 28/29c and 30d/31a Conodont at Shangsi. The two earliest Triassic size reduction episodes were correlative between the two sections. These Hindeodus-Isarcicella changes coincide with some important geological events such as eustatic sea-level changes, anoxic events, carbon Size reduction isotope oscillations, miniaturization of brachiopods and microbial changes. Through detailed investigation of the Permian–Triassic transition palaeoenvironment and the palaeoecology of Hindeodus–Isarcicella, the authors propose that the main causes of South China the size reduction was a sharp decline of food availability because of the mass extinction and the anoxic event during the P/T transition. The pattern of size reduction supports suggestions that the end-Permian mass extinction was multi-episodal, consisting of 3 extinction events rather than a single catastrophic event. © 2008 Elsevier B.V. All rights reserved. 1. Introduction There are several opinions about the causes and patterns of the P–T mass extinction (Isozaki, 1997; Kozur, 1998; Wang and Cao, 2004; The end-Permian biotic crisis was the largest mass extinction in Fang, 2004a,b; Grice et al., 2005; Racki and Wignall, 2005). Some the fossil record. It eliminated over 90% of species in the oceans authors have proposed a single-episode catastrophic mass extinction (Stanley and Yang, 1994; Bambach et al., 2004) and about 70% of (Jin et al., 2000; Kaiho et al., 2006), while others have argued for a vertebrate families on land (Benton, 1988; King, 1991; Maxwell, 1992). multi-episode mass extinction (Wu and Liu, 1991; Wignall and The cause or causes and duration as well as the nature of the Hallam, 1993; Yin and Tong, 1998; Fang, 2004a,b; Xie et al., 2005; extinction remain uncertain and actively debated (Wu and Liu, 1991; Shen et al., 2006; Yin et al., 2007a). In this paper, we attempt to test Wignall and Hallam, 1993; Isozaki, 1997; Kozur, 1998; Yin and Tong, these various scenarios by using the size variation data of a group of 1998; Jin et al., 2000; Yin et al., 2001; Wang and Cao, 2004; Fang, conodont species from a single clade from several continuous 2004a,b; Grice et al., 2005; Racki and Wignall, 2005; Yin et al., 2007a). Permian–Triassic boundary sections in South China. The fundamental The Global Stratotype Section and Point (GSSP) of the Permian– question addressed in the study is to see if the sizes of conodont Triassic Boundary at Meishan in Zhejiang Province, China has served species varied across the PTB and, if so, whether or not the timing of as a focal point in this global debate, as it has provided much critical the significant size changes actually corresponded to any of the stratigraphical and palaeontological data. As a result, the section has proposed PTB extinction intervals. A related question, also investi- received intensive multidisciplinary studies by various research gated as an integral part of this study, is to elucidate the possible cause groups, including lithostratigraphy, biostratigraphy, sedimentology, (s) for the size change of the conodont species across the PTB. sequence stratigraphy, isotope geochemistry, eventostratigraphy, and There is now a considerable literature relating size variation in magnetostratigraphy (Yin et al., 2001 and references therein). lineages through time to biotic crises caused by environmental changes in earth history. Initially, Urbanek (1993) coined the term “Lilliput effect” for an observed size reduction of Silurian graptolites ⁎ Corresponding author. Faculty of Earth Sciences, China University of Geosciences, Wuhan, Hubei 430074, PR China. Tel.: +86 27 67883139; fax: +86 27 87515956. during a biotic crisis. Subsequently, other researchers have reported E-mail address: [email protected] (X. Lai). similar size decreases during times of extinction; for example, the size 0031-0182/$ – see front matter © 2008 Elsevier B.V. All rights reserved. doi:10.1016/j.palaeo.2008.04.015 G. Luo et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 264 (2008) 176–187 177 Fig. 1. A: Location map of study area in South China; Meishan Section (B) and Shangsi Section (C). (after Wingnall et al., 1995 and Jiang et al., 2007). of late Devonian conodonts (Girard and Renaud, 1996), heart urchins study of large conodont samples. Furthermore, unlike most other across the Cretaceous/Tertiary boundary (Jeffery, 2001), and various organisms, most Late Permian conodont species or their lineages organisms around the Permian/Triassic boundary (Twitchett, 2001; persisted through into the Lower Triassic. Therefore, these conodonts He et al., 2005; Twitchett, 2005a,b; Luo et al., 2006; Twitchett, 2006; can serve as the best material for the study of the size variation during He et al., 2007; Twitchett, 2007). the P–T transitional period. Luo et al. (2006) reported a size reduction It is widely held that there was no obvious change in conodont in P1 elements of the conodont genus Neogondolella at the bed 24d/ fortunes during the end-Permian mass extinction, because many 24e junction (Upper Permian) at Meishan. However, the limited conodont lineages clearly survived through the PTB (e. g. Clark et al., Neogondolella specimens from the Lower Triassic did not allow us to 1986; Jiang et al., 2007). However, the survival of lineages tends to perform a full-scale size variation study throughout the Permian– overlook the potential ecological information that can come from the Triassic transition. To overcome this shortfall, in this paper we have Table 1 The total number, mean size, standard deviation, percentage of specimens larger than 0.5 mm and 95% confidence interval of the mean for the P1 element of Hindeodus–Isarcicella from the P/T transition at the Meishan Section, Changxing, Zhejiang Province Bed 24a 24b 24c 24d 24e 25 26 27a 27b 27c 27d 28 29 Number 79 39 5 77 17 2 8 94 133 141 270 171 47 Mean (mm) 0.458 0.459 0.443 0.534 0.436 0.533 0.392 0.455 0.503 0.421 0.467 0.580 0.459 Standard deviation 0.132 0.143 0.079 0.096 0.100 0.006 0.043 0.098 0.108 0.103 0.158 0.150 0.108 Percentage (N0.5mm) 29.11 33.33 20.00 76.62 25.53 100 0 27.66 47.37 18.44 34.44 67.84 38.30 95% confidence interval of the mean 0.4913 0.4989 0.7002 0.5610 0.4951 0.5838 0.4048 0.4725 0.5182 0.4372 0.4871 0.6008 0.4926 0.4244 0.4026 0.1831 0.5144 0.3595 0.4822 0.3619 0.4298 0.4811 0.4017 0.4472 0.5536 0.4264 178 G. Luo et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 264 (2008) 176–187 chosen hindeodid conodonts because as a lineage they survived across only previously been undertaken for the P –T transition (Luo et al., the P–T boundary. The recovery and size measurement of abundant 2006). The present paper examines the size variations, in large conodont specimens is time-consuming, and this kind of study has samples, of conodonts from the Hindeodus–Isarcicella clade during the Fig. 2. Size distribution histogram of P1 elements of Hindeodus–Isarcicella from each bed in ascending order (24a to 29 except for bed 25) through the P/T transition at the Meishan Section A. The figures above each histogram are the total number of specimens in each size range. G. Luo et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 264 (2008) 176–187 179 end Permian to earliest Triassic interval, with the aim of improving our tions of the evolution of the hindeodid–isarcicellid lineage are understanding of the mass extinction patterns and of biotic recovery. controversial (Kozur, 1996; Ding et al., 1997; Lai, 1997; Wang and Wang, 1997). Ding et al. (1997) thought that Hindeodus latidentatus–H. 2. Material and method parvus–Isarcicella turgida–I. isarcica formed an anagenetic lineage. However, Wang and Wang (1997) replaced I. turgida with I. staeschei Thirteen successive bulk samples ranging from bed 24 to bed 29 in the evolutionary sequence of (Ding et al., 1997). Nevertheless, (the PTB is sited at the bottom of bed 27c) were collected from Section almost all authors have accepted the idea of an evolutionary sequence A at Meishan, Changxing, Zhejiang province (Fig.
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