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Implications for the Evolution of Early Danian Planktonic Foraminifera

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Implications for the Evolution of Early Danian Planktonic Foraminifera Accepted Manuscript Revalidation of the genus Chiloguembelitria Hofker: Implications for the evolution of early Danian planktonic foraminifera Ignacio Arenillas, José A. Arz, Vicente Gilabert PII: S1464-343X(17)30297-2 DOI: 10.1016/j.jafrearsci.2017.07.011 Reference: AES 2963 To appear in: Journal of African Earth Sciences Received Date: 24 August 2016 Revised Date: 15 May 2017 Accepted Date: 17 July 2017 Please cite this article as: Arenillas, I., Arz, José.A., Gilabert, V., Revalidation of the genus Chiloguembelitria Hofker: Implications for the evolution of early Danian planktonic foraminifera, Journal of African Earth Sciences (2017), doi: 10.1016/j.jafrearsci.2017.07.011. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. ACCEPTED MANUSCRIPT 1 Revalidation of the genus Chiloguembelitria Hofker: implications for 2 the evolution of early Danian planktonic foraminifera 3 4 Ignacio Arenillas*, José A. Arz and Vicente Gilabert 5 6 Address of the authors: 7 * Corresponding author at: Departamento de Ciencias de la Tierra, and Instituto 8 Universitario de Investigación en Ciencias Ambientales de Aragón, Universidad de 9 Zaragoza, E-50009 Zaragoza, Spain. Tel: +34976762475. E-mail address: 10 [email protected] 11 12 José A. Arz: Departamento de Ciencias de la Tierra, and Instituto Universitario de 13 Investigación en Ciencias Ambientales de Aragón, Universidad de Zaragoza, E-50009 14 Zaragoza, Spain. Tel: +34976762507. E-mail address: [email protected] 15 16 Vicente Gilabert: Departamento de Ciencias de la Tierra, Universidad de Zaragoza, E- 17 50009 Zaragoza, Spain. Tel: +34660286216. E-mail address: [email protected] 18 19 1 ACCEPTED MANUSCRIPT 20 ABSTRACT 21 Guembelitria is the only planktonic foraminiferal genus whose survival from the 22 mass extinction event of the Cretaceous/Paleogene (K/Pg) boundary has been clearly 23 proven. The evolution of Guembelitria after the K/Pg boundary led to the appearance of 24 two guembelitriid lineages in the early Danian: one biserial, represented by 25 Woodringina and culminating in Chiloguembelina, and the other trochospiral, 26 represented by Trochoguembelitria and culminating in Globoconusa. We have re- 27 examined the genus Chiloguembelitria, another guembelitriid descended from 28 Guembelitria and whose taxonomic validity had been questioned, it being considered a 29 junior synonym of the latter. Nevertheless, Chiloguembelitria differs from Guembelitria 30 mainly in the wall texture (pustulate to rugose vs. pore-mounded) and the position of the 31 aperture (umbilical-extraumbilical to extraumbilical vs. umbilical). Chiloguembelitria 32 shares its wall texture with Trochoguembelitria and some of the earliest specimens of 33 Woodringina, suggesting that it played an important role in the evolution of early 34 Danian guembelitriids, as it seems to be the most immediate ancestor of both 35 trochospiral and biserial lineages. Morphological and morphostatistical analyses of 36 Chiloguembelitria discriminate at least five species: Chg. danica, Chg. irregularis, and 37 three new species: Chg. hofkeri, Chg. trilobata and Chg. biseriata. 38 39 Keywords: Guembelitriids, wall texture, K/Pg boundary, morphostatistical analysis, 40 Tunisia. 41 42 1. Introduction 43 The mass extinction event of the Cretaceous/Paleogene (K/Pg) boundary 66 million 44 years ago eliminated almost all species of Maastrichtian planktonic foraminifera (Smit 2 ACCEPTED MANUSCRIPT 45 1990; Arenillas et al. 2002; Molina et al. 2006, 2009), leaving vacant most of the 46 pelagic niches and triggering in the early Danian the most important radiation in their 47 evolutionary history. One of the most passionate debates in the Earth Sciences focuses 48 on the paleobiological and paleoenvironmental changes that occurred before, during and 49 after this extinction, as well as its relation with the massive eruptions in the Deccan 50 volcanic province in India (Chenet et al. 2007; Schoene et al. 2015) and/or with the 51 Chicxulub asteroid impact on Yucatan in Mexico (Hildebrand et al. 1991; Schulte et al. 52 2010). 53 Numerous new species of trochospiral and biserial planktonic foraminifera originated 54 after the K/Pg boundary (Luterbacher and Premoli Silva 1964; Smit 1982; Canudo et al. 55 1991; Liu and Olsson 1992; Molina et al. 1998). This evolutionary radiation happened 56 in two pulses (Arenillas et al. 2000b, 2004). The first occurred between approximately 5 57 and 20 kyr after the K/Pg boundary (Arenillas et al. 2016b), with the appearance of 58 species belonging to the parvularugoglobigerinids (Parvularugoglobigerina Hofker, 59 1978, and Palaeoglobigerina Arenillas, Arz and Náñez, 2007) and biserial taxa 60 (Woodringina Loeblich and Tappan, 1957, and Chiloguembelina Loeblich and Tappan, 61 1956). The second evolutionary radiation occurred between approximately 37 and 80 62 kyr after the K/Pg boundary, giving rise to species belonging to Trochoguembelitria 63 Arenillas, Arz and Náñez, 2012, Eoglobigerina Morozova, 1959, Parasubbotina 64 Olsson, Berggren and Liu, 1992, Globanomalina Haque, 1956, and Praemurica Olsson, 65 Hemleben, Berggren and Liu 1992 (Arenillas et al. 2010, 2012; Arenillas and Arz 66 2013a, 2013b, 2016a). Other genera appear shortly afterwards, such as Subbotina 67 Brotzen and Pozaryska, 1961, and Globoconusa Khalilov, 1956. 68 One of the presumed ancestors of the earliest Danian taxa was Guembelitria 69 Cushman, 1933, the only planktonic foraminiferal genus whose survival from the K/Pg 3 ACCEPTED MANUSCRIPT 70 mass extinction event has been clearly proven (Smit 1982; Olsson et al. 1999; Arenillas 71 et al. 2000a; Ashckenazi-Polidova et al. 2014; Arenillas et al. 2016a). There is a general 72 consensus that Guembelitria is the ancestor of microperforate genera such as 73 Woodringina and Globoconusa (Olsson et al. 1999; Arenillas and Arz 2000; Arenillas et 74 al. 2010; Koutsoukos 2014). For the latter, Arenillas et al. (2012, 2016b) proposed the 75 evolutionary lineage Guembelitria-Trochoguembelitria-Globoconusa, instead of the 76 more direct derivation of Globoconusa from Guembelitria. Woodringina, with a mixed 77 triserial-biserial test, is in turn the ancestor of the wholly biserial genus 78 Chiloguembelina. 79 Guembelitria species were r-strategy opportunists that inhabited surface-water 80 environments (Nederbragt 1991) and bloomed during the stressful times of 81 Maastrichtian global warming events associated with the Deccan Traps eruptions (Pardo 82 and Keller 2008). Guembelitria also bloomed immediately after the Chicxulub impact, 83 during approximately the first 10 or 15 kyr of the Danian (acme-stage PFAS-1 of 84 Arenillas et al. 2006). Another, later bloom of triserial guembelitriids has been 85 recognized in the early Danian of Egypt, Israel, Tunisia and India. This was related to a 86 global warming episode linked to the last phase of Deccan volcanism (Punekar et al. 87 2014). 88 The main object of the present study is Chiloguembelitria Hofker, 1978, another 89 guembelitriid that originated in the first evolutionary radiation and whose taxonomic 90 validity has been questioned, it being considered a junior synonym of Guembelitria (e.g. 91 D'Hondt 1991; MacLeod 1993). However, Chiloguembelitria may be key to elucidating 92 the evolutionary relationships among the earliest Danian guembelitriids. Arenillas et al. 93 (2010) suggested that Chiloguembelitria includes at least three species: Chg. danica 94 Hofker, 1978, Chg. irregularis (Morozova, 1961) and Ch. cf. cretacea. However, 4 ACCEPTED MANUSCRIPT 95 studies of its morphologic variability and species diversity have not been conducted so far. 96 far. 97 In this paper, we document new specimens assignable to the genus 98 Chiloguembelitria mainly from the El Kef section (Tunisia) in order to assess its 99 taxonomic validity, advance the understanding of its phylogenetic relationships with 100 Guembelitria, Woodringina and other genera, and determine its species diversity. This 101 review will also help to date and correlate the climatic warming episodes of the early 102 Danian. The bloom of triserial guembelitriids linked to the last volcanic phase of the 103 Deccan has been ascribed to Guembelitria (Punekar et al. 2014). Nevertheless, it could 104 in fact be an acme of Chiloguembelitria, which replaced Guembelitria in the early 105 Danian, occupying the same ecological niche. Considering Chiloguembelitria and 106 Guembelitria as separate genera will make it possible to differentiate more easily the 107 possible Danian blooms of Chiloguembelitria from the PFAS-1 episode (acme of 108 Guembelitria inmediately after the K/Pg boundary), recognize and calibrate possible 109 hiatuses in lower Danian sections, and interpret and correlate more accurately the 110 paleoenvironmental changes occurring after the K/Pg boundary extinction event. 111 112 2. Material and Methods 113 Samples for this study were selected from the lower Danian of the El Kef section, 114 Tunisia, which is the Global boundary Stratotype Section and Point for the base of the 115 Danian Stage (Molina et al. 2006). All studied rock samples were disaggregated in 116 water with diluted
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