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Early Triassic Disaster and Opportunistic Foraminifers in South China

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Early Triassic Disaster and Opportunistic Foraminifers in South China This is a repository copy of Early Triassic disaster and opportunistic foraminifers in South China. White Rose Research Online URL for this paper: http://eprints.whiterose.ac.uk/90242/ Version: Accepted Version Article: Song, H, Tong, J, Wignall, PB et al. (5 more authors) (2016) Early Triassic disaster and opportunistic foraminifers in South China. Geological Magazine, 153 (2). pp. 298-315. ISSN 0016-7568 https://doi.org/10.1017/S0016756815000497 Reuse Unless indicated otherwise, fulltext items are protected by copyright with all rights reserved. The copyright exception in section 29 of the Copyright, Designs and Patents Act 1988 allows the making of a single copy solely for the purpose of non-commercial research or private study within the limits of fair dealing. The publisher or other rights-holder may allow further reproduction and re-use of this version - refer to the White Rose Research Online record for this item. Where records identify the publisher as the copyright holder, users can verify any specific terms of use on the publisher’s website. Takedown If you consider content in White Rose Research Online to be in breach of UK law, please notify us by emailing [email protected] including the URL of the record and the reason for the withdrawal request. [email protected] https://eprints.whiterose.ac.uk/ 1 Early Triassic disaster and opportunistic 2 foraminifers in South China 3 Haijun Song*à , Jinnan Tong* , Paul B. Wignall§, 4 Mao Luo||, Li Tian*, Huyue Song*, YunFei Huang¶, Daoliang Chu* 5 *State Key Laboratory of Biogeology and Environmental Geology, China University of Geosciences, 6 Wuhan 430074, PR China 7 àState Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and 8 Palaeontology, Nanjing, 210008, PR China 9 §School of Earth and Environment, University of Leeds, Leeds LS2 9JT, UK 10 ||School of Life and Environmental Sciences, Deakin University, Melbourne Burwood Campus, 11 Burwood, Victoria 3125, Australia 12 ¶School of Geoscience, Yangtze University, Wuhan 430100, PR China 13 Author for correspondence: [email protected] (H. Song), [email protected] (J. Tong) 14 15 Abstract: Survival and recovery are important dynamic processes of biotic evolution 16 during major geological transitions. Disaster and opportunistic taxa are two significant 17 groups that dominate the ecosystem in the aftermath of mass extinction events. Disaster 18 taxa appear immediately after such crises whilst opportunists predate the crisis but also 19 bloom in the aftermath. This paper documents three disaster foraminiferal species and 20 seven opportunistic foraminiferal species from Lower Triassic successions of South China. 21 They are characterized by extreme high-abundance and low-diversity and occurred 22 occasionally in the Griesbachian, Smithian, and Spathian. The characteristics (small size, 23 simple morphology) and stratigraphic ranges of these groups suggest that r-selection is a 24 commonly used strategy for survivors to cope with either harsh post-extinction conditions 25 and/or environments lacking incumbents. 26 Keywords: disaster, opportunist, foraminifers, Early Triassic, Permian-Triassic extinction 27 28 1. Introduction 29 For the survivors of mass extinctions, their fate can be highly variable but also to some 30 extent predictable. Groups with intrinsically high rates of extinction before the crisis often 31 radiate at high rates afterwards whilst evolutionary laggards often recover much more 32 slowly. This is exemplified by the ammonoids, a group characterized by exceptionally high 33 evolutionary rates throughout their history. Having suffered a severe extinction at the 34 Permian-Triassic (P-Tr) they radiated at typically high rates in the immediate aftermath 35 (Stanley, 2007). Similarly, the bivalves, the evolutionary carthorses of the marine 36 invertebrates, underwent little radiation in the aftermath of the P-Tr extinction with the 37 exception of the spectacular recovery of the claraiids, a family of “flat clams” belonging to 38 the Pterinopectinidae, that exhibit similarly high evolutionary rates before the extinction 39 (Yin, 1985). 40 One of the most interesting facets of the immediate post-extinction interval is the 41 presence of prolific abundances of opportunists, typically called disaster taxa (Harries, 42 Kauffman & Hansen, 1996; Kauffman & Harries, 1996). These are defined as species that 43 are adapted to the high stress conditions of an extinction crisis and its immediate aftermath 44 but are rare or absent at other times (Harries, Kauffman & Hansen, 1996; Kauffman & 45 Harries, 1996; Rodland & Bottjer, 2001). They thus differ from “normal” opportunists, 46 which have long species ranges, and often appear in high stress settings including (but not 47 restricted to) the aftermath of mass extinctions. Opportunists are, by definition, ecological 48 generalists that exhibit high fecundity – a facet of their lifestyle that is manifest by the 49 rapid attainment of sexual maturity and a small, simple morphology. Disaster taxa also 50 typically exhibit these features but their more restricted temporal distribution suggests that 51 they are suited to the specific and unusual conditions of the post-extinction interval. 52 Disaster taxa are evolutionary dead ends. However, they differ from a third category found 53 at this time – progenitor taxa which appear and radiate rapidly in the post-extinction 54 interval (Kauffman & Harries, 1996). 55 Several benthic biota have been considered as potential disaster forms in the aftermath 56 of the P-Tr extinction e.g. stromatolites (Schubert & Bottjer, 1992), lingulide brachiopods 57 (Rodland & Bottjer, 2001) and calcareous tubeworms (He et al., 2012). Amongst the 58 foraminifers the small, tube-like Earlandia is regarded as a typical disaster genus found in 59 huge numbers immediately following both the Frasnian-Famennian (F-F) mass extinction 60 and the Permian-Triassic boundary (PTB) extinctions (Hallam & Wignall, 1997). 61 Earlandia is known in the aftermath of both P-Tr extinction pulses (Song et al., 2013) 62 along with Postcladella kahlori. For example both these taxa are especially abundant in the 63 microbialite facies that developed following the latest Permian extinction in Turkey 64 (Altiner et al., 1980; Altiner & Zaninetti, 1981; Groves, Altiner & Rettori, 2005), Italy 65 (Groves et al., 2007) and South China (Song et al., 2009). Earlandia also occurs in 66 wackestones the immediate aftermath of the earliest Triassic extinction at Meishan section 67 of South China (Wignall & Twitchett, 2002). 68 However, the ecological significance of disaster taxa is unclear. Traditionally, 69 opportunists should record high-stress environmental conditions and so, ostensibly, the 70 presence of post-extinction disaster forms could record the persistence of high-stress 71 conditions that caused the preceding mass extinction. Alternatively they may record the 72 expansion of hardy opportunists, capable of surviving the extinction episode, into vacated 73 environments once the environments had returned to normal. In this second alternative 74 disaster taxa fit a distinct ecological category (rather than a temporally-defined subset of 75 opportunist) – their success is due to their extinction-resistance but not to any specific 76 adaptation to the environments in which they find themselves in the post-extinction world. 77 The significance of disaster taxa is at the heart of a long-running debate on the delayed 78 recovery of benthic ecosystems in the Early Triassic in the aftermath of the P-Tr mass 79 extinction. Hallam (1991) was the first to note that the severity of the P-Tr mass extinction 80 and delayed recovery may be, in part, due to the prolongation of the harmful conditions that 81 triggered the extinction – specifically the extent and duration of global marine anoxia. In 82 contrast, Schubert & Bottjer (1992; 1995) noting the spread of stromatolites in Early 83 Triassic seas argued that they were filling an ecospace in which biotic factors (such as 84 gastropod grazing) were much reduced (and had yet to recover) but with normal, physical 85 environmental factors. Similarly, Rodland & Bottjer’s (2001) work on lingulide 86 brachiopods in the Early Triassic of the western USA concluded that their proliferation 87 took place in well-oxygenated shelf seas. In contrast, Pruss & Bottjer (2004) and Fraiser & 88 Bottjer (2009) studied impoverished Early Triassic trace fossil assemblages in the same 89 strata and suggested repetition of stressful conditions. In support of this conclusion, 90 contemporaneous trace fossils from nearer shore strata in western Canada are much higher 91 diversity suggesting that there was indeed something stressful about offshore marine 92 settings in the Early Triassic (Zonneveld et al., 2010). 93 In this study, we document the types, stratigraphic ranges and ecological behaviors of 94 disaster and opportunistic foraminifers during the biotic recovery from the P-Tr mass 95 extinction and address the issue of whether they were survivors living in a pleasant but 96 emptied nirvana or whether they were living in a harsh post-apocalyptic hell. 97 98 2. Geological setting and studied sections 99 We report on our analysis of the foraminifer content of the Lower Triassic successions 100 of South China and supplement our observations with literature records from elsewhere. 101 During the P-Tr transition, the South China block was located in the eastern Tethys near 102 the equator, consisting of islands,
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