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A Highly Diverse Bivalve Fauna from a Bithynian (Anisian, Middle Triassic) Tubiphytes-Microbial Buildup in North Dobrogea (Romania)

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A Highly Diverse Bivalve Fauna from a Bithynian (Anisian, Middle Triassic) Tubiphytes-Microbial Buildup in North Dobrogea (Romania) Zurich Open Repository and Archive University of Zurich Main Library Strickhofstrasse 39 CH-8057 Zurich www.zora.uzh.ch Year: 2021 A highly diverse bivalve fauna from a Bithynian (Anisian, Middle Triassic) Tubiphytes ‐microbial buildup in North Dobrogea (Romania) Friesenbichler, Evelyn ; Hautmann, Michael ; Grădinaru, Eugen ; Bucher, Hugo Abstract: This paper describes a rich bivalve fauna from the Tubiphytes‐Limestone Member (Bithynian, early middle Anisian) of the Caerace Formation in North Dobrogea (Romania). We report 51 bivalve species, including 7 new species and 2 new genera: Atrina multicostata sp. nov., Pinna simionescui sp. nov., Chlamys (Praechlamys) prima sp. nov., Entolium reticulatum sp. nov., Scythentolium anisicum sp. nov., Romaniamya mahmudiaensis gen. et sp. nov. and Praedicerocardium vetulus gen. et sp. nov. The latter taxon represents the oldest Triassic megalodontoid. We provide detailed descriptions of the bivalve fauna, discuss the ecology and compare species richness in the Early and Middle Triassic, showing that the bivalve assemblage of North Dobrogea is much more diverse than all previously described Early Triassic bivalve faunas, but not as rich as several faunas from the late Anisian and Ladinian. The relative richness of the Dobrogea fauna with respect to geologically older and younger faunas suggests that it represents an early stage of the main rediversification of bivalves after the end‐Permian mass extinction. DOI: https://doi.org/10.1002/spp2.1286 Posted at the Zurich Open Repository and Archive, University of Zurich ZORA URL: https://doi.org/10.5167/uzh-175886 Journal Article Accepted Version Originally published at: Friesenbichler, Evelyn; Hautmann, Michael; Grădinaru, Eugen; Bucher, Hugo (2021). A highly diverse bivalve fauna from a Bithynian (Anisian, Middle Triassic) Tubiphytes ‐microbial buildup in North Do- brogea (Romania). Papers in Palaeontology, 7(1):447-495. DOI: https://doi.org/10.1002/spp2.1286 Palaeontology A highly diverse bivalve fauna from a Bithynian (Anisian, Middle Triassic) Tubiphytes-microbial buildup in North Dobrogea (Romania) Journal: Palaeontology Manuscript ID PALA-10-18-4331-OA.R2 Manuscript Type: Original Article Date Submitted by the n/a Author: Complete List of Authors: Friesenbichler, Evelyn; Universitat Zurich Palaontologisches Institut und Museum, Hautmann, Michael; University Zurich, Palaeontologisches Institut und Museum Grădinaru, Eugen; Universitatea din Bucuresti, Department of Geology Bucher, Hugo; Paläontologisches Institut und Museum, Universität Zürich; Key words: Bivalvia, Anisian, Bithynian, North Dobrogea, diversity, recovery Palaeontology Page 1 of 145 Palaeontology 1 2 3 4 5 1 A highly diverse bivalve fauna from a Bithynian (Anisian, Middle Triassic) Tubiphytes-microbial 6 7 2 buildup in North Dobrogea (Romania) 8 9 10 3 11 12 1 1 2 1 13 4 Evelyn Friesenbichler *, Michael Hautmann , Eugen Grădinaru , Hugo Bucher , 14 15 16 5 17 18 19 6 1 Paläontologisches Institut und Museum, University of Zurich, Karl-Schmid-Strasse 4, 8006 Zurich, 20 21 7 Switzerland; e-mails: [email protected], [email protected], 22 23 8 [email protected] 24 25 26 9 2 Department of Geology, Faculty of Geology & Geophysics, University of Bucharest, Bd. Bălcescu 27 28 29 10 Nicolae 1, RO-010041, Bucharest, Romania; e-mail: [email protected] 30 31 32 11 33 34 35 12 *corresponding author 36 37 38 13 39 40 41 14 42 43 44 45 15 Abstract 46 47 16 48 49 50 17 This paper describes a rich bivalve fauna from the Tubiphytes-Limestone Member (Bithynian, early 51 52 53 18 middle Anisian) of the Caerace Formation in North Dobrogea (Romania). We report 51 bivalve 54 55 19 species, including seven new species and two new genera: Atrina multicostata n. sp., Pinna 56 57 20 simionescui n. sp., Chlamys (Praechlamys) prima n. sp., Entolium reticulatum n. sp., Scythentolium 58 59 21 anisicum n. sp., Romaniamya mahmudiaensis n. gen. et n. sp. and Praedicerocardium vetulus n. gen. 60 1 Palaeontology Palaeontology Page 2 of 145 1 2 3 22 et n. sp. The latter taxon represents the oldest Triassic megalodontoid. We provide detailed 4 5 23 descriptions of the bivalve fauna, discuss the ecology and compare species richness in the Early and 6 7 8 24 Middle Triassic, showing that the bivalve assemblage of North Dobrogea is much more diverse than 9 10 25 all previously described Early Triassic bivalve faunas, but not as rich as several faunas from the late 11 12 26 Anisian and Ladinian. The relative richness of the Dobrogea fauna with respect to geologically older 13 14 27 and younger faunas suggests that it represents an early stage of the main rediversification of bivalves 15 16 17 28 after the end-Permian mass extinction. 18 19 20 29 21 22 23 30 24 25 26 27 31 Keywords: Bivalvia, Anisian, Bithynian, North Dobrogea, diversity, recovery. 28 29 32 30 31 32 33 33 34 35 34 The most severe biotic crisis in Earth’s history, the end-Permian mass extinction about 252 Ma ago 36 37 38 35 (Burgess et al. 2014), did not only extinguish between 81% (Stanley 2016) and 96% (Raup 1979) of all 39 40 36 marine species, but also caused a temporal shift from the Upper Paleozoic T-type (‘tropical’) 41 42 37 carbonate factory to a non-skeletal M-type (‘mud-mound’) carbonate factory (sensu Schlager 2000, 43 44 38 2003; Baud & Richoz 2004; Baud et al. 2007). While M-factory reefs already flourished during the 45 46 39 Early Triassic (e.g. Brayard et al. 2011; Friesenbichler et al. 2018), T-factory reefs started 47 48 49 40 re-diversifying since the Middle Triassic (Flügel & Stanley 1984; Stanley 1988; Senowbari-Daryan et 50 51 41 al. 1993; Flügel 1994, 2002; Martindale et al. 2019). One of the last ‘pure’ M-type Triassic carbonate 52 53 42 factories before the transition toward T-type factories is a Tubiphytes-microbial buildup located in 54 55 43 the Mahmudia Quarry in the North Dobrogea Orogen (Popa et al. 2014), which is also one of the few 56 57 58 44 bioconstructions from the Tethyan realm that flourished during the Bithynian (early middle Anisian, 59 60 45 Middle Triassic). 2 Palaeontology Page 3 of 145 Palaeontology 1 2 3 46 The in situ boundstones of the Tubiphytes-microbial buildup developed on the upper slope of a distal 4 5 47 platform in relatively deep water under normal oxic conditions and in an intertropical environment 6 7 8 48 (Forel & Grădinaru 2018). It was built up by a micro-framework of Tubiphytes, a small encrusting and 9 10 49 low-tiering organism of uncertain affinities (Riding & Guo 1992) that is a common fossil in Anisian 11 12 50 reef assemblages. Apart from the micro-framework of Tubiphytes, the buildup contains abundant 13 14 51 fine-grained internal sediments, microbial crusts and autochthonous micrite, added by a large 15 16 17 52 volume of marine cements (Popa et al. 2014). The Tubiphytes-microbial buildup provided a suitable 18 19 53 environment for various groups of abundant and diverse benthic macroinvertebrates such as 20 21 54 bivalves, gastropods (Nützel et al. 2018) and brachiopods (Grădinaru & Gaetani 2019), plus a rich 22 23 55 ostracod fauna (Forel & Grădinaru 2018). 24 25 26 56 The Bithynian Tubiphytes-microbial buildup is one of the oldest Triassic carbonate reefs and provides 27 28 29 57 the opportunity to study how bivalves adapted to this new habitat type and how this affected their 30 31 58 recovery after the end-Permian mass extinction. This study is part of a series of studies (Forel & 32 33 59 Grădinaru 2018; Nützel et al. 2018; Grădinaru & Gaetani 2019) that aim at the systematic description 34 35 60 and interpretation of the fossil content from the Tubiphytes-microbial buildup in North Dobrogea. 36 37 38 61 We report 51 bivalve species belonging to 39 genera, which makes bivalves the most diverse 39 40 62 macroinvertebrate group of the Tubiphytes-microbial buildup so far. Seven species and two genera 41 42 63 are newly described, including the oldest Triassic representative of the Megalodontoida. The ecology 43 44 64 of the bivalve fauna is reconstructed and its species richness is discussed in the context of the 45 46 65 rediversification of marine life after the end-Permian mass extinction. 47 48 49 50 66 51 52 53 67 54 55 56 57 68 GEOLOGICAL AND STRATIGRAPHICAL SETTING 58 59 69 60 3 Palaeontology Palaeontology Page 4 of 145 1 2 3 70 The North Dobrogea Orogen is located in the foreland of the Alpine Carpathian Orogen (e.g. 4 5 71 Săndulescu 1995) and is limited by the Danube Delta in the North and the Black Sea in the East. It is 6 7 8 72 the westernmost segment of the Palaeo-Tethyan Cimmeride Orogenic System and continues 9 10 73 eastward into the Mountainous Crimea, the North Caucasus (Fig. 1A, inset map) and further 11 12 74 eastwards into the Asian Cimmerides (Săndulescu 1980; Şengör 1984, 1986). The North Dobrogea 13 14 75 Orogen is a fold-and-thrust belt that consists of several tectonic units (Fig. 1A. Visarion et al. 1990; 15 16 17 76 Săndulescu 1995). The Triassic succession has its widest extent In the Tulcea Unit, which is the 18 19 77 lowermost tectonic unit of the North Dobrogea Orogen and is located in its eastern part. The Triassic 20 21 78 sediments unconformably overlie the Variscan basement and are developed both in basinal and 22 23 79 carbonate platform facies (Grădinaru 1995, 2000). During the Middle Triassic, a carbonate platform 24 25 80 developed in the central and eastern areas of the Tulcea Unit. The various carbonate facies that 26 27 28 81 developed in the eastern area of the Tulcea Unit are described in detail by Popa et al.
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