Zurich Open Repository and Archive University of Zurich Main Library Strickhofstrasse 39 CH-8057 Zurich www.zora.uzh.ch

Year: 2019

Palaeoecology of Late Ladinian (Middle Triassic) benthic faunas from the Schlern/Sciliar and Seiser Alm/Alpe di Siusi area (South Tyrol, Italy)

Friesenbichler, Evelyn ; Hautmann, Michael ; Nützel, Alexander ; Urlichs, Max ; Bucher, Hugo

Abstract: The Schlern and Seiser Alm area (South Tyrol, Italy) is a classical locality for studies of Middle Triassic platform to basin transitions, yet details of the palaeoecology of the rich benthic faunas of this area have been insufficiently known. We present herein a detailed palaeoecological study of thefauna from the Schlernplateau beds (Late Ladinian to Early Carnian) and the more or less time-equivalent Pachycardientuffe (Late Ladinian), which is based on quantitative faunal data. Both the palaeoecology and sedimentary features suggest that the fauna of the Schlernplateau beds represents a lagoonal soft- bottom fauna. The high species richness of the fauna and the locally restricted occurrences of fossils indicate an open-lagoon setting palaeogeographically close to an ocean inlet. The high evenness of the fauna is probably a result of time-averaging. In contrast, the fauna of the Pachycardientuffe shows clear indications of transportation. Ecological features of this fauna andpalaeogeographic reconstructions suggest three potential source areas: (1) the lagoon represented by the Schlernplateau beds, (2) the reef fringing this lagoon and (3) a shallow clastic coast of a nearby volcanic island and/or submarine high.A comparison between diversities of selected Early and Middle Triassic lithological units revealed the increasing species richness of all major benthic taxa during the Middle Triassic and a shift from bivalve-dominated Early Triassic faunas to gastropod-dominated faunas. Keywords Middle Triassic · Biotic recovery · Quantitative faunal analyses · Palaeoecology · Palaeoenvironment

DOI: https://doi.org/10.1007/s12542-018-0423-7

Posted at the Zurich Open Repository and Archive, University of Zurich ZORA URL: https://doi.org/10.5167/uzh-152680 Journal Article Accepted Version

Originally published at: Friesenbichler, Evelyn; Hautmann, Michael; Nützel, Alexander; Urlichs, Max; Bucher, Hugo (2019). Palaeoecology of Late Ladinian (Middle Triassic) benthic faunas from the Schlern/Sciliar and Seiser Alm/Alpe di Siusi area (South Tyrol, Italy). Paläontologische Zeitschrift, 93(1):1-29. DOI: https://doi.org/10.1007/s12542-018-0423-7 Pa lZ Palaeoecology of Late Ladinian (Middle Triassic) benthic faunas from the Schlern/Sciliar and Seiser Alm/Alpe di Siusi area (South Tyrol, Italy) --Manuscript Draft--

Manuscript Number: PAZE-D-17-00052R2 Full Title: Palaeoecology of Late Ladinian (Middle Triassic) benthic faunas from the Schlern/Sciliar and Seiser Alm/Alpe di Siusi area (South Tyrol, Italy) Article Type: Research Paper Corresponding Author: Evelyn Friesenbichler, MSc Universitat Zurich Zurich, SWITZERLAND Corresponding Author Secondary Information: Corresponding Author's Institution: Universitat Zurich Corresponding Author's Secondary Institution: First Author: Evelyn Friesenbichler, MSc First Author Secondary Information: Order of Authors: Evelyn Friesenbichler, MSc Michael Hautmann Alexander Nützel Max Urlichs Hugo Bucher Order of Authors Secondary Information:

Funding Information: Schweizerischer Nationalfonds zur Dr. Michael Hautmann Förderung der Wissenschaftlichen Forschung (200021-162402/1)

Abstract: The Schlern and Seiser Alm area (South Tyrol, Italy) is a classical locality for studies of Middle Triassic platform to basin transitions, yet details of the palaeoecology of the rich benthic faunas of this area have been insufficiently known. We herein present a detailed palaeoecological study of the fauna from the Schlernplateau beds (late Ladinian to early Carnian) and the more or less time-equivalent Pachycardientuffe (late Ladinian), which is based on quantitative faunal data. The palaeoecology as well as sedimentary features suggest that the fauna of the Schlernplateau beds represents a lagoonal soft bottom fauna. The high species richness of the fauna as well as the locally restricted occurrences of fossils indicate an open lagoonal setting palaeogeographically close to an oceanic inlet. The high evenness of the fauna is probably a result of time-averaging. In contrast, the fauna of the Pachycardientuffe shows clear indications of transportation. Ecological features of this fauna and palaeogeographic reconstructions suggest three potential source areas: (1) the lagoon represented by the Schlernplateau beds, (2) the reef fringing this lagoon and (3) a shallow clastic coast of a nearby volcanic island and/or submarine high. A comparison between diversities of selected Early and Middle Triassic lithological units revealed the increasing species richness of all major benthic taxa during the Middle Triassic and a shift from bivalve dominated Early Triassic faunas to gastropod dominated faunas. Response to Reviewers: Ref.: Ms. No. PAZE-D-17-00052R1 Palaeoecology of Late Ladinian (Middle Triassic) benthic faunas from the Schlern/Sciliar and Seiser Alm/Alpe di Siusi area (South Tyrol, Italy)

Powered by Editorial Manager® and ProduXion Manager® from Aries Systems Corporation General note: Please see attached document with “track changes” for the details of our changes in response to the reviewer’s comments.

Reviewer’s comments

AUTHOR AFFILIATIONS: Once again, to avoid any confusion, please please split the affiliations of Alexander Nützel ... SNSB-BSPG and LMU are clearly different institutions (see my former comments)

Response: We split the affiliation of Alexander Nützel and apologize that we did not do that in the first place.

TEXT: Page 22, line 505 ... Szulc INSTEAD OF Sculz 2007 Page 23, line 524 ... Kozur and Mostler 1973 IS MISSING in your REFERENCES Page 23, lines 524/525 ... Mostler 1977 IS MISSING in your REFERENCES Page 23, line 526 ... Reich 2013 IS MISSING in your REFERENCES Page 24, line 548 ... Gaetani et al. 1981 INSTEAD OF Gaetani 1981 Page 28, line 646 ... Eichwald, E. von. 1851. INSTEAD OF Eichwald, M. 1851. Page 28, line 646 ... Geognosie, auf INSTEAD OF Geognosie auf Page 28, line 647 ... die Eifel, Tyrol, Italien, Sizilien INSTEAD OF die Eife, Tirol, Sizilien Page 30, line 691 ... Gümbel, C.W. 1861. INSTEAD OF Gümbel, C.W. von 1861. ...(Gümbel was nobled in 1882) Page 33, lines 754/55 ... implement Kozur and Mostler 1973 (see Page 23, line 524) Page 34, lines 782/783 ... implement Mostler 1977 (see Page 23, lines 524/525) Page 36, lines 820/821 ... implement Reich 2013 (see Page 23, line 526) Page 36, lines 825-826 ... DELETE Sano and Nakashima 1997 which is not cited in text etc. Page 39, line 887 ... "Trachyceras" IN ITALICS please Page 39, line 889 ... "Myophoria kefersteini" IN ITALICS please Page 40, line 928 ... Yin INSTEAD OF Ying Page 41, line 930 ... Yin INSTEAD OF Ying Page 41, line 932 ... Yin INSTEAD OF Ying Page 43 ... line 981 ... von Eichwald, 1851 INSTEAD OF Eichwald, 1851

Response: As requested we changed the above mentioned text passages and deleted the work of Sano and Nakazawa (1997). Naturally, we added the work of Kozur and Mostler (1973), Mostler (1977) and Reich (2013) to the reference list and apologize that this mistake did not track our attention before.

REFERENCES: Please check again my former comments (comment 5) ... Please use an en-dash NOT a hyphen netween page numbers

Response: All hyphens between page numbers were changed into en-dashes.

TABLE: Frizzell and Exline (1956) INSTEAD OF Frizzell and Exline (1955)

Response: We changed the above mentioned text passage as requested.

ONLINE RESOURCE: Please re-format your species names ... "sp.", "sp. 1...", "cf.", "n.sp." NOT IN ITALICS

Response: We performed the necessary changes in our electronic supplementary material.

Powered by Editorial Manager® and ProduXion Manager® from Aries Systems Corporation Further changes

Independently from the reviewer’s suggestions we performed some minor changes on the manuscript, which can be seen in the track-edited version of the manuscript.

Powered by Editorial Manager® and ProduXion Manager® from Aries Systems Corporation Manuscript Click here to download Manuscript Manuscript revision 2.docx

Click here to view linked References

1 2 1 Palaeoecology of Late Ladinian (Middle Triassic) benthic faunas from the Schlern/Sciliar and Seiser 3 4 2 Alm/Alpe di Siusi area (South Tyrol, Italy) 5 6 7 3 8 9 10 4 Evelyn Friesenbichler1*, Michael Hautmann1, Alexander Nützel2, Max Urlichs3, Hugo Bucher1 11 12 13 5 14 15 16 1 17 6 Paläontologisches Institut und Museum, University of Zurich, Karl-Schmid-Strasse 4, 8006 Zurich, 18 19 7 Switzerland 20 21 22 8 2 SNSB - Bayerische Staatssammlung für Paläontologie und Geologie, Richard-Wagner-Straße 10, 23 24 25 9 80333 Munich, Germany and Department of Earth and Environmental Sciences, Palaeontology & 26 27 10 Geobiology, Geo-Bio-Center LMU, Richard-Wagner-Straße 10, 80333 Munich, Germany 28 29 30 11 3 Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany 31 32 33 12 34 35 36 37 13 * corresponding author; e-mail: [email protected]; tel: +41 44 634 23 47 38 39 40 14 41 42 43 44 15 Acknowledgements 45 46 47 16 48 49 50 17 This study was supported by the Swiss National Science Foundation project 200021-162402/1 (to 51 52 18 M. Hautmann). Markus Hebeisen is thanked for his support during the preparation of fossil material 53 54 55 19 and Rosi Roth is thanked for helping during photographing the fossils. Michael Rasser (State Museum 56 57 20 of Natural History Stuttgart, Germany) kindly provided access to the collection of the Stuttgart State 58 59 60 21 Museum of Natural History and lending some specimens of the Stuttgart collection. This article 61 62 63 1 64 65 22 benefited from careful reviews by William Foster, Christopher A. McRoberts, Simon Schneider and 1 2 23 Mike Reich. 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 2 64 65 24 Abstract 1 2 25 3 4 5 6 26 The Schlern and Seiser Alm area (South Tyrol, Italy) is a classical locality for studies of Middle 7 8 27 Triassic platform to basin transitions, yet details of the palaeoecology of the rich benthic faunas of 9 10 28 this area have been insufficiently known. We herein present a detailed palaeoecological study of the 11 12 13 29 fauna from the Schlernplateau beds (late Ladinian to early Carnian) and the more or less time- 14 15 30 equivalent Pachycardientuffe (late Ladinian), which is based on quantitative faunal data. The 16 17 31 palaeoecology as well as sedimentary features suggest that the fauna of the Schlernplateau beds 18 19 20 32 represents a lagoonal soft bottom fauna. The high species richness of the fauna as well as the locally 21 22 33 restricted occurrences of fossils indicate an open lagoonal setting palaeogeographically close to an 23 24 25 34 oceanic inlet. The high evenness of the fauna is probably a result of time-averaging. In contrast, the 26 27 35 fauna of the Pachycardientuffe shows clear indications of transportation. Ecological features of this 28 29 36 fauna and palaeogeographic reconstructions suggest three potential source areas: (1) the lagoon 30 31 32 37 represented by the Schlernplateau beds, (2) the reef fringing this lagoon and (3) a shallow clastic 33 34 38 coast of a nearby volcanic island and/or submarine high. A comparison between diversities of 35 36 39 selected Early and Middle Triassic lithological units revealed the increasing species richness of all 37 38 39 40 major benthic taxa during the Middle Triassic and a shift from bivalve dominated Early Triassic faunas 40 41 41 to gastropod dominated faunas. 42 43 44 42 45 46 47 48 43 Keywords: Middle Triassic, biotic recovery, quantitative faunal analyses, palaeoecology, 49 50 44 palaeoenvironment. 51 52 53 45 54 55 56 57 58 46 Kurzfassung 59 60 47 61 62 63 3 64 65 48 Obwohl das Gebiet des Schlern und der Seiser Alm (Südtirol, Italien) seit langem als klassische 1 2 49 Lokalität für die Erforschung mitteltriassischer Plattform-Becken-Übergänge gilt, ist die Paläoökologie 3 4 5 50 ihrer reichen benthischen Fauna unzureichend erforscht. Wir präsentieren eine detaillierte 6 7 51 paläoökologische Studie der Fauna der Schlernplateau-Schichten (spätes Ladinium bis frühes 8 9 52 Karnium) und der mehr oder weniger gleichalten Pachycardientuffe (spätes Ladinium), welche auf 10 11 12 53 quantitativen Daten basiert. Die Paläoökologie sowie der sedimentologische Kontext legen nahe, 13 14 54 dass die Fauna der Schlernplateau-Schichten eine lagunäre Weichbodenfauna repräsentiert. Ihre 15 16 17 55 hohe Diversität sowie die lokal begrenzten Vorkommen der Fossilien deuten auf eine offene Lagune 18 19 56 hin, die sich paläogeographisch nahe an einem Einlass zum offenen Meer befand. Die hohe 20 21 57 Gleichmäßigkeit der Fauna ist höchstwahrscheinlich das Resultat von „time-averaging“. Im Gegensatz 22 23 24 58 dazu gibt es in der Fauna der Pachycardientuffe klare Hinweise auf Transport. Die Ökologie dieser 25 26 59 Fauna und paläogeographische Rekonstruktionen lassen auf drei potentielle Herkunftsgebiete 27 28 60 schließen: (1) die Lagune, welche durch die Schlernplateau-Schichten repräsentiert wird, (2) das 29 30 31 61 Saumriff dieser Lagune und (3) eine flache, siliziklastische Küste nahe einer vulkanischen Insel 32 33 62 und/oder einer submarinen Hochzone. Ein Vergleich der Diversitäten ausgewählter früh- und 34 35 63 mitteltriassischer lithologischer Einheiten offenbart einen Anstieg der Diversität während der Früh- 36 37 38 64 und Mitteltrias sowie einen Wechsel von Muschel-dominierten frühtriassischen Faunen zu 39 40 65 Gastropoden-dominierten Faunen während der Mitteltrias. 41 42 43 44 66 45 46 47 67 Schlüsselwörter: Mitteltrias, ökologische Erholung, quantitative Faunen-Analysen, Paläoökologie, 48 49 68 Paläo-Umwelt. 50 51 52 69 53 54 55 56 57 70 1. Introduction 58 59 71 60 61 62 63 4 64 65 72 The recovery from the end-Permian mass extinction, which killed between 81% (Stanley 2016) 1 2 73 and 96% (Raup 1979) of marine species, provides a unique test-case for studying long-term 3 4 5 74 evolutionary responses to a sudden and massive loss of species on a global scale. Scientific interest in 6 7 75 studying the recovery pattern is twofold: it allows testing macroevolutionary models (e.g., Brayard et 8 9 76 al. 2009; Hautmann et al. 2015), and it represents a (pessimistic) analogue for the long-term future of 10 11 12 77 biosphere in the face of anthropogenically caused environmental changes (Payne and Clapham 13 14 78 2012). 15 16 17 79 Understanding evolutionary processes requires the study of species in their ecological context. 18 19 20 80 Concerning the recovery from the end-Permian mass extinction, much palaeoecological work has 21 22 81 been done in the past decades, but these studies almost exclusively concentrated on the Early 23 24 25 82 Triassic (e.g., Schubert and Bottjer 1995; Twitchett and Wignall 1996; Hofmann et al. 2013a, 2013b, 26 27 83 2014, 2015; Hautmann et al. 2011, 2015; Foster et al. 2017), which is the ca. 4 Ma time interval that 28 29 84 followed the end-Permian mass extinction. However, biodiversity in terms of global taxonomic 30 31 32 85 richness did not reach the pre-extinction level before the Late Jurassic (Sepkoski 1997) or later (Alroy 33 34 86 et al. 2008). With regard to alpha-diversity in local environments, the major recovery step seems to 35 36 87 have taken place during the Middle Triassic, as evidenced by the striking differences between most 37 38 39 88 diverse Early Triassic communities that include around 45 benthic species (Hautmann et al. 2013, 40 41 89 2015; Hofmann et al. 2013b, 2014) and, e.g., the hyperdiversified fauna from the Longobardian– 42 43 90 Julian (Late Ladinian Early Carnian) Cassian Fm. of northern Italy (>1000 benthic species; Tab. 1). The 44 – 45 46 91 progress in recovery during the Middle Triassic has been addressed in several recent studies (e.g., 47 48 92 Payne 2005; Payne et al. 2006; Song et al. 2011; Velledits et al. 2011; Foster and Sebe 2017) but data 49 50 51 93 are still relatively scarce. 52 53 54 94 This study is based on quantitative bed-rock sampling in the area of Schlern/Sciliar mountain and 55 56 95 Seiser Alm/Alpe di Siusi (South Tyrol, Italy; Fig. 1), which is a paradigm for a Middle to Late Triassic 57 58 59 96 reef-to-basin landscape. It is one of the few places in the world where the platform-slope-basin 60 61 97 transitional zone is preserved in its primary setting (Keim 2008; Brandner et al. 2016), giving the 62 63 5 64 65 98 opportunity to study the ecology of concurrent faunas in different environments. Sampling was 1 2 99 carried out (1) in sediments of a lagoonal setting represented by the locally highly fossiliferous 3 4 5 100 Schlernplateau beds on the Schlern mountain and (2) in the basinal setting of the ‘Pachycardientuffe’ 6 7 101 at the Seiser Alm, which represent approximately the same stratigraphic age (see chapter 3). The 8 9 102 palaeoecology and species distributions of both faunas are discussed in the context of their 10 11 12 103 depositional environments, and their diversities are compared with published data on Early–Middle 13 14 104 Triassic benthic communities from other regions, which allows preliminary insights into diversity 15 16 17 105 changes through the Early and Middle Triassic time interval. 18 19 20 106 21 22 23 24 107 2. Material and Methods 25 26 27 108 28 29 30 109 Fieldwork was carried out in August 2016 at the Seiser Alm/Alpe di Siusi and the Schlern/Sciliar 31 32 110 mountain in South Tyrol/Alto Adige in northern Italy. At the Seiser Alm, fossils were sampled in the 33 34 35 111 gorge of the Frommerbach/Rio Frommer and one of its eastern tributaries (Figs 1 and 2a-b). On the 36 37 112 Schlern, fossils were mainly collected on the western flank of the Seiser Klamm/Gola di Siusi (Figs 1 38 39 113 and 2c-d). Outcropping rock material of fossiliferous beds was quantitatively sampled until the 40 41 42 114 occurrences of fossils were exhausted. Additionally, fossils quantitatively collected by M. Urlichs 43 44 115 between 1998 and 2012 at the same localities were included in the analyses. Specimens used for the 45 46 116 palaeoecological analyses are housed at the State Museum of Natural History Stuttgart (Germany, 47 48 49 117 inventory numbers SMNHS17501/250 - SMNHS75538) and the Museum of Nature South Tyrol (Italy, 50 51 118 inventory numbers PZO5980 - PZO5998). For taxonomic identification fossils were first washed and 52 53 54 119 then, when necessary, prepared with micro jacks. A quantitative matrix of fossil occurrences was 55 56 120 compiled at species level (Online Resource 1). For bivalves, the number of individuals has been 57 58 121 inferred by counting only the most numerous valves (left or right) together with articulated 59 60 61 122 specimens. Only fossils that (1) were clearly relatable to specific beds of the investigated outcrops 62 63 6 64 65 123 and (2) samples with more than 40 individuals were used for palaeoecological data analyses. 1 2 124 Sedimentological observations in the field and information from ecological analyses were used for 3 4 5 125 the interpretation of the depositional environment. Our complete dataset comprises 2723 individuals 6 7 126 assigned to 53 species for the Schlernplateau beds and 651 individuals assigned to 59 species for the 8 9 127 Pachycardientuffe. 10 11 12 13 128 Ecological parameters and statistical analyses were calculated using the software PAST, version 14 15 129 2.17 (Hammer et al. 2001). Based on the species-abundance data, rarefaction analyses were carried 16 17 130 out for each sample in order to compare species richness of faunas with different sample sizes and to 18 19 20 131 assess the relative completeness of sampling. For further ecological characterization, rank- 21 22 132 abundance distributions, trophic nuclei, and ecological pie charts were used. The trophic nucleus of 23 24 25 133 an association comprises those species that contribute 80% of the total number of individuals in an 26 27 134 association (Neyman 1967). Following Aberhan (1994), species were assigned to 10 trophic guilds, 28 29 135 based on functional morphology, mode of life of related Recent species, information from the 30 31 32 136 literature, and the sedimentological context. Gastropods were collectively assigned to epifaunal 33 34 137 herbivores and/or detritus-feeders, since evidence for more specialized feeding lacks. For the 35 36 138 identification of recurrent assemblages and groups of ecologically related species, Q-mode cluster 37 38 39 139 analyses were calculated for samples containing more than 40 individuals, using the Morisita 40 41 140 similarity index (Morisita 1959) and unweighted pair-group average. Additionally, R-mode cluster 42 43 44 141 analyses were used for the identification of sets of species that typically co-occur. 45 46 47 142 48 49 50 51 143 3. Geologic setting 52 53 54 144 55 56 57 145 The Schlern and the adjacent Rosengarten/Catinaccio represent a Middle Triassic carbonate 58 59 146 platform with northeastward, eastward to southeastward inclined reef slopes (Brandner 1991; 60 61 62 63 7 64 65 147 Bosellini 1998; Brandner et al. 2007; Keim 2008). Towards the west, there was probably a connection 1 2 148 to the Mendel platform (Keim 2008). The Schlern Mountain is build up by (1) the pre-volcanic, up to 3 4 5 149 800 m thick ‘Schlerndolomite’ (Rosengarten Fm.), (2) up to 40 m thick volcanic rocks and (3) post- 6 7 150 volcanic, well-bedded dolomite formerly known as ‘Rosetta Dolomite’ or ‘Cassian Dolomite’, now 8 9 151 interpreted as the platform facies of the Rosszähne Fm. The latter is overlain by (4) the 10 11 12 152 Schlernplateau beds, which is a sequence of carbonate rocks and grey to reddish marls that are 13 14 153 locally highly fossiliferous and commonly interpreted as lagoonal sediments (Figs 2c-d; Keim 2008; 15 16 17 154 Brandner et al. 2016). Findings of the biostratigraphically important ammonoid Prototrachyceras 18 19 155 archelaus (Laube, 1869b) indicate a Late Ladinian age of the Schlernplateau beds (Brandner 1991; 20 21 156 Urlichs and Tichy 2000). However, the occurrence of the bivalve Myophoria kefersteini kefersteini 22 23 24 157 (Münster in Goldfuss, 1837) in the uppermost 6 m of the Schlernplateau beds suggests that the top 25 26 158 of the formation extends into the Lower Carnian (Urlichs 2014). The thickness of the Schlernplateau 27 28 159 beds varies from only a few meters to a maximum of ca. 40 m. Towards the platform margins, the 29 30 31 160 Schlernplateau beds interfinger with the reef flat facies. 32 33 34 161 During deposition of the Rosszähne Fm. and Schlernplateau beds at the top of the Schlern 35 36 162 carbonate platform, the adjacent Seiser Alm basin was filled with a more than 500 m thick 37 38 39 163 successions of volcanoclastic detritus of the Wengen Fm. and, later, the mixed siliciclastic-carbonate 40 41 164 succession of the St. Cassian/San Cassiano Fm. (Brandner 1991). Within the Wengen Fm. of the Seiser 42 43 165 Alm, the named for the mass 44 Late Ladinian (Urlichs 1977, 1994, 2014) ‘Pachycardientuffe’, which are 45 46 166 occurrence of the bivalve Pachycardia rugosa von Hauer, 1857, represent several (at least two) 47 48 167 conglomeratic, highly fossiliferous intercalations that have been mapped as distal parts of the 49 50 51 168 ‘Marmolada Conglomerate’ in the geological map of the western Dolomites (Autonome Provinz 52 53 169 Bozen (ed.) 2007). However, the Marmolada Conglomerate in the strict sense is a volcanogenic 54 55 170 conglomerate that was transported from a source area near the Marmolada Mountain (ca. 20 km 56 57 58 171 east of the study area) into adjacent basins (e.g., Bosellini 1998; Bottoli and Trombetta 1998). In 59 60 172 contrast, the lithology of the Pachycardientuffe is dominated by non-dolomitized limestone pebbles 61 62 63 8 64 65 173 and cobbles (small to medium sized ‘Cipit boulders’) that originate from adjacent carbonate 1 2 174 platforms. We therefore assume that the Pachycardientuffe represent distal equivalents of tongues 3 4 5 175 of foreslope talus deposits, as described by Yose (1991) in a proximal facies at the Mahlknecht cliff, 6 7 176 around 4 km SE of our sampling localities. The admixed volcanogenic pebbles and volcanoclastic 8 9 177 matrix could either have been taken up during transportation, or stem from a second source area, or 10 11 12 178 could represent the result of mixing of carbonate and volcanogenic components by reworking (see 13 14 179 chapter 5.2.). 15 16 17 180 18 19 20 21 22 181 4. The Schlernplateau beds 23 24 182 25 26 27 28 183 Early workers erroneously correlated the fauna of the Schlernplateau beds with the younger Raibl 29 30 184 Fm. (Carnian), based among other reasons on the occurrence of Myophoria kefersteini (Münster in 31 32 185 Goldfuss, 1837; e.g., von Richthofen 1860). This view was also shared by von Wöhrmann and Koken 33 34 35 186 (1892) in their comprehensive description of the Schlernplateau fauna and was maintained by most 36 37 187 subsequent authors until Brandner (1991), Urlichs and Tichy (2000) and Urlichs (2014) provided 38 39 188 evidence for a Late Ladinian (Prototrachyceras archelarus Frankites regoledanus ammonoid zone) to 40 – 41 42 189 early Carnian (Daxatina canadensis ammonoid zone) age. In addition to ammonoid finds, the key 43 44 190 observation for the revised stratigraphy was the existence of several subspecies of Myophoria 45 46 191 kefersteini, which are indicative of different stratigraphic intervals (Urlichs 2014). With this updated 47 48 49 192 stratigraphic information, Schlernplateau beds and Pachycardientuffe are more or less time- 50 51 193 equivalent units. 52 53 54 194 Von Wöhrmann and Koken (1892) described 72 species, including 33 gastropod species, 24 55 56 57 195 bivalve species, eight cephalopod species, three echinoid species, two coral species and one crinoid 58 59 196 species from the Schlernplateau beds. Notably, nearly all specimens of this rich fauna stem from a 60 61 62 63 9 64 65 197 limited area at the western flank and southern part of the Seiser Klamm. The only other locality that 1 2 198 is explicitly mentioned in von Wöhrmann and Koken (1892) is the ‘Burgstall’, some hundred meters 3 4 5 199 east of the Seiser Klamm, where one coral species (Thecosmilia rothpletzi von Wöhrmann and Koken, 6 7 200 1892), fragments of crinoids and echinoid spines were found. Our survey confirms that fossils are 8 9 201 scarce or absent at other places of the Schlernplateau. At the Seiser Klamm locality, fossils are most 10 11 12 202 abundant in the upper part of the section, particularly in several layers of red marl (Fig. 3). Fossils are 13 14 203 also abundant in several rudstone beds near the top of the section (Fig. 2e), but the crystallized 15 16 17 204 limestone makes it difficult to extract them from the matrix. 18 19 20 205 A detailed section was measured at the western flank of the Seiser Klamm and samples were 21 22 206 taken from three localities on the Maultierrücken/Dorsale del Mul (Figs 1 and 3). The investigated 23 24 25 207 section starts with an alternation (ca. 20 m) of partly dolomitic limestone beds, which are separated 26 27 208 by marls or covered intervals that probably also represent fine-grained sediments. The limestone and 28 29 209 dolomite beds are mud-, wacke-, pack- and grainstones that sometimes contain bioclasts like bivalve 30 31 32 210 and gastropod shells and, less common, ooids. The overlaying marl beds (SP3 and SP5) are highly 33 34 211 fossiliferous. Above these marls, an around 3 m thick sequence of thick-bedded wackestone forms a 35 36 212 prominent cliff, the top of which is formed by a 30 cm thick oolitic limestone bed (at a height of 25 37 38 39 213 m). Several marly, highly fossiliferous intervals above this cliff provided the majority of fossils studied 40 41 214 herein. Within the marly intervals, laterally discontinuous oolithic limestone beds are repeatedly 42 43 215 intercalated, which sometimes contain bioclasts such as gastropod and bivalve shells. A notable 44 45 46 216 lithological marker bed near the top of the section contains shell debris, ooids, and interspersed 47 48 217 pisolithic iron ore grains. Surface ornamentations on the fossils are generally well preserved. This lack 49 50 51 218 of abrasion as well as the fact that some bivalves were preserved with conjoined valves indicate that 52 53 219 the fossils were not transported over larger distances. According to Urlichs and Tichy (2000) and 54 55 220 Urlichs (2014) the uppermost layers of the investigated section are already Carnian in age. 56 57 58 221 Accordingly, the Ladinian/Carnian boundary lies within fossiliferous bed SP10 (Fig. 3). 59 60 61 222 62 63 10 64 65 223 4.1. Fauna of the Schlernplateau beds 1 2 224 3 4 5 6 225 A total number of 53 species occurring in 16 fossiliferous beds (SP1-SP16, Fig. 3) was identified, 7 8 226 including 27 gastropod species, 19 bivalve species, four coral species and one sponge species (Online 9 10 227 Resource 1). Notably, brachiopods are absent, and only one crinoid columnal fragment and one 11 12 13 228 echinoid spine were found. In terms of species richness, our collection is comparable to the diversity 14 15 229 described in the monograph of von Wöhrmann and Koken (1892; 72 species). Species richness within 16 17 230 the different beds varies between 2 to 32. However, rarefaction analyses of samples from each fossil 18 19 20 231 horizon indicate different degrees of sampling completeness (Fig. 4a). 21 22 23 232 The 10 most frequent species are, in descending order, Myophoria kefersteini (Münster in 24 25 233 Goldfuss, 1837) (bivalve, 16.3%), Heminajas fissidentata (von Wöhrmann, 1889) (bivalve, 11.4%), 26 27 28 234 Pachycardia rugosa von Hauer, 1857 (bivalve, 9.7%), Neritaria plicatilis (von Klipstein, 1844) 29 30 235 (gastropod, 8.3%), Naticopsis neritacea (Münster, 1841) (gastropod, 8%), Coelostylina solida (von 31 32 33 236 Wöhrmann and Koken, 1892) (gastropod, 5.8%), Palaeonarica concentrica (Münster, 1841) 34 35 237 (gastropod, 4.3%), Myophoriopis richthofeni (Stur, 1868) (bivalve, 4%), Trigonodus rablensis (Gredler, 36 37 238 1862) (bivalve, 3.8%) and ‘Loxonema’ aequale von Wöhrmann and Koken, 1892 (gastropod, 3.3%). 38 39 40 239 These 10 species together with Cassianella bipartita (von Klipstein, 1845) (bivalve, 3.2%) and 41 42 240 Stephanocosmia coronata (von Wöhrmann and Koken, 1892) (gastropod, 3%; Fig. 5 and 6) constitute 43 44 241 the trophic nucleus of the complete fauna at the Schlernplateau. Simpson’s index of dominance of 45 46 47 242 associations from different fossiliferous beds is generally high and varies between 0.06 and 0.30, with 48 49 243 an average value of 0.2 (Online Resource 1). 50 51 52 244 Cluster analysis (Fig. 7) did not reveal the existence of clearly distinct associations. However, a 53 54 55 245 cluster of samples that are dominated by Myophoria kefersteini is set off from samples in which 56 57 246 species of Pachycardia and Trigonodus are more abundant. Because Myophoria, Trigonodus and 58 59 247 Pachycardia are not only phylogenetically closely related but also ecologically similar (both are 60 61 62 63 11 64 65 248 shallow infaunal suspension feeders), we do not believe that these two clusters represent 1 2 249 fundamentally different environmental conditions. As an alternative explanation, it seems possible 3 4 5 250 that haphazardness of larvae settlement had randomly favoured different dominant species at 6 7 251 different times. 8 9 10 252 Although the fauna of the Schlernplateau beds is gastropod dominated in terms of species 11 12 13 253 richness, it is bivalve dominated in terms of total number of individuals. Accordingly, most species 14 15 254 were epifaunal herbivores and/or detritus-feeders, whereas almost half of all collected individuals 16 17 255 are shallow infaunal suspension-feeders, followed by epifaunal herbivores and/or detritus-feeders 18 19 20 256 and free-lying epifaunal suspension-feeders (Fig. 8a). However, the guild-specific diversity and 21 22 257 absolute abundance of trophic guilds can vary between different fossiliferous beds (Fig. 9). 23 24 25 258 Frequency distributions of assemblages from different fossiliferous beds show different patterns, 26 27 28 259 but a remarkable aspect is that many show an unusually high degree of evenness (Figs 9b-c). The lack 29 30 260 of abrasion, occurrence of bivalves with conjoined valves, and the sedimentological context 31 32 33 261 (occurrence in fine-grained sediments) exclude that this pattern is due to effects of faunal mixing by 34 35 262 transportation. Another notable aspect is that there are often lateral differences in the faunal 36 37 263 composition within single fossiliferous beds (Fig. 9). 38 39 40 41 264 42 43 44 265 4.2. Interpretation of the Schlernplateau fauna 45 46 266 47 48 49 267 The geologic context leaves little doubt that the Schlernplateau beds represent lagoonal 50 51 52 268 sediments of the platform interior. This is evident by the sedimentary features of the Schlernplateau 53 54 269 beds themselves, which include the coexistence of high- and low-energy sediments (e.g., oolites and 55 56 57 270 shell coquinas (rudstones) versus marls, wackestones, and early diagenetic dolomites) and the 58 59 271 presence of locally abundant oncoids (Fig. 2f). It is further corroborated by the lateral facies change, 60 61 272 which includes the transition to fringing coral reefs on the supposed seaward side and to pedogenic 62 63 12 64 65 273 sediments at places where the platform was emerged (Brandner et al. 2016 and own observations). 1 2 274 Finally, the ecology of the documented fauna is in accordance with a lagoonal setting because 3 4 5 275 shallow infaunal suspension-feeders and free-lying epifaunal suspension-feeders dominate, which 6 7 276 are commonly associated with a soft substratum. 8 9 10 277 Given the lagoonal setting, two aspects of the Schlernplateau fauna are remarkable. First, the 11 12 13 278 species richness is unusually high for this environment. Although decidedly stenohaline organisms 14 15 279 lack (e.g., brachiopods) or are extremely rare (e.g., echinoderms and ammonoids), the high species 16 17 280 richness excludes that the lagoon was (at the place where the samples were taken) affected by high 18 19 20 281 environmental stress such as changing salinity, which often characterizes restricted lagoons (e.g., 21 22 282 Fürsich 1994; Fürsich et al. 1995). The second unusual aspect is the extremely high evenness in many 23 24 25 283 samples. Based on the state of preservation and the occurrence in marly beds (see above), faunal 26 27 284 mixing by transportation can be excluded. However, no examples of primary causes for such high 28 29 285 evenness are known to us from ecologically comparable faunas. We therefore assume that the high 30 31 32 286 evenness may, at least partly, reflect the effects of time-averaging. Fürsich and Aberhan (1990) nicely 33 34 287 demonstrated how time-averaging may produce abundance patterns similar to those that we 35 36 288 observed in the Schlernplateau beds. 37 38 39 40 289 Our summary interpretation of the sedimentological and faunistic data is that the Schlernplateau 41 42 290 beds represent an open lagoon and that the outcrop area around the Seiser Klamm was 43 44 291 palaeogeographically close to an oceanic inlet that prevented large-scale fluctuations in salinity and 45 46 47 292 temperature, provided supply of food for filter feeding organisms, and was the source of larvae that 48 49 293 settled within the lagoon. The randomness in the composition of larvae that reached the lagoon may 50 51 294 be partly responsible for differences in the faunal composition. In this model, successive 52 53 54 295 communities went back to different settlement events, each with a particular taxonomic composition 55 56 296 of larvae and thus resulting in different dominance of the resulting cohort. Superimposition of 57 58 59 297 abundance patterns of successive communities by time-averaging then caused a pattern where many 60 61 298 species are nearly equally abundant. However, prevalent species probably dominated at different 62 63 13 64 65 299 times. We note that time-averaging might have also contributed to the high overall species richness, 1 2 300 but even if this effect is considered, the Schlernplateau beds are still remarkably species rich. 3 4 5 6 301 Additional support for the open lagoon model comes from the fact that fossil occurrences are 7 8 302 locally restricted (close to oceanic inlets in our interpretation). From a sedimentological perspective, 9 10 303 the laterally discontinuous oolitic limestone beds that are intercalated in the low-energy marls 11 12 13 304 probably represent washover deposits from a nearby barrier, which however was discontinuous near 14 15 305 the fossil locality. 16 17 18 306 19 20 21 22 23 307 5. The Pachycardientuffe/Tufi a Pachycardie 24 25 308 26 27 28 309 Like the fauna from the Schlernplateau beds, the highly diverse fauna of the Pachycardientuffe is 29 30 31 310 known for more than 100 years. Broili (1903) described 157 species, including 122 bivalve species, 18 32 33 311 brachiopod species, 13 echinoid species and four crinoid species. Blaschke (1905) added the 34 35 36 312 description of 77 gastropod species, as well as three sponge species, five coral species, one 37 38 313 scaphopod species and 14 bivalve species that were not described in Broili (1903). A detailed 39 40 314 description of brachiopods from the Pachycardientuffe was given by Waagen (1903). After a revision 41 42 43 315 by Waagen (1907) a total number of 165 bivalve species is described from the Pachycardientuffe. 44 45 316 Overall, more than 270 species have been reported from the Pachycardientuffe thus far (Tab. 1). It 46 47 317 should be noted, however, that some of the major fossil localities (e.g., at the Tschapitbach/Rio Cipit) 48 49 50 318 are currently covered and therefore not accessible. 51 52 53 319 The investigated section at locality Frommerbach A is located in the gorge of the Frommerbach 54 55 320 itself (Figs 1, 2a and 10). It starts with a fossiliferous conglomerate at 1850 m altitude that overlies 56 57 58 321 volcanoclastic sandstone beds of the Wengen Fm. with an erosional disconformity. The pebbles are 59 60 322 mainly limestones and less commonly rounded volcanic rocks, which are embedded in a siltstone 61 62 63 14 64 65 323 matrix. The overlaying bedded sandstone contains limestone blocks at its top. The components of 1 2 324 the following fossiliferous conglomerate consist of limestone and volcanoclastics. After a 3 m thick 3 4 5 325 interval of siltstone, highly fossiliferous sandy siltstones follow on top of this section. The 6 7 326 investigated section Frommerbach B is located in an eastern tributary of the Frommerbach (Figs 1, 2b 8 9 327 and 10) at 1850 m altitude. The rocks are matrix supported, fossiliferous, pebble bearing, 10 11 12 328 volcanogenic, middle- to coarse grained sandstones. As pointed out by Urlichs (2014), the 13 14 329 fossiliferous rocks in the two Frommerbach sections do not belong to the same horizon. The 15 16 17 330 investigated rocks of section Frommerbach A are younger than the rocks from Frommerbach B. For a 18 19 331 detailed stratigraphic correlation between the sections see Fig. 3 in Urlichs (2014). 20 21 22 332 23 24 25 333 5.1. Fauna of the Pachycardientuffe 26 27 28 334 29 30 31 335 At the two Frommerbach localities a total number of 59 species was found, which is much less 32 33 336 than the overall species richness of the Pachycardientuffe (277 species), as described in Broili (1903), 34 35 36 337 Waagen (1903), Blaschke (1905) and Waagen (1907). These 59 species include 30 bivalve species, 16 37 38 338 gastropod species, five echinoid species, three coral species, three crinoid species and two 39 40 41 339 brachiopod species. Sponges are absent. The seven most frequent species are, in descending order, 42 43 340 Pachycardia rugosa von Hauer, 1857 (54.4%), Pachycardia plieningeri Broili, 1903 (27.3%), 44 45 341 Trigonodus rablensis (Gredler, 1862) (3.7%), Myophoria kefersteini (Münster in Goldfuss, 1837) 46 47 48 342 (1.8%), Fedaiella inaequiplicata (von Klipstein, 1843) (gastropod, 1.4%), Naticopsis seisiensis 49 50 343 (Blaschke, 1905) (gastropod 1.2%) and Heminajas fissidentata (von Wöhrmann, 1889) (1.1%; Figs 5 51 52 344 and 6). The fauna is clearly dominated by the two bivalve species Pachycardia rugosa and 53 54 55 345 Pachycardia plieningeri that form the trophic nucleus. All other species are relatively rare and almost 56 57 346 half of the species is only represented by one individual (Figs 8b and 11). Simpson-D ranges between 58 59 60 61 62 63 15 64 65 347 0.29 and 0.47, with an average value of 0.39 (Online Resource 1). Again, the rarefaction analysis 1 2 348 shows that the diversity of some samples is underestimated (Fig. 4b). 3 4 5 6 349 A palaeontological analysis of the fauna from the Pachycardientuffe was already performed by 7 8 350 Fürsich and Wendt (1977), based on 283 individuals belonging to 26 species, including bivalves (11 9 10 351 species), gastropods (11 species), crinoids (two species), echinoids and corals (one species, 11 12 13 352 respectively). This analysis shows a similar rank-abundance pattern as in our study. Pachycardia 14 15 353 rugosa is with 85.5% the most abundant species, whereas almost three-quarters of all species are 16 17 354 represented by only one individual. Because Fürsich and Wendt (1977) did not separate Pachycardia 18 19 20 355 rugosa and Pachycardia plieningeri, the Simpson-D in their study is relatively higher (0.71) than in our 21 22 356 study (min. D = 0.29; max. D = 0.46). Whether P. rugosa and P. plieningeri represent two different 23 24 25 357 species is debatable. In this study, we used the less deeply depressed lunule and the more posterior 26 27 358 position of the beaks in P. plieningeri as criteria for their separation. Additional criteria noted by 28 29 359 Broili (1903) include a more elongated outline and more elongated, but narrower anterior cardinal 30 31 32 360 tooth in P. plieningeri. 33 34 35 361 The fauna of the Pachycardientuffe is bivalve dominated in terms of species richness and total 36 37 362 number of individuals. The guild-species diversity shows that epifaunal herbivores and/or detritus- 38 39 40 363 feeders, epibyssate suspension-feeders and shallow infaunal suspension-feeders are the most 41 42 364 important guilds (Fig. 8b). However, most individuals belong to shallow to moderate deep infaunal 43 44 365 suspension-feeders, which is not surprising since both dominating species belong to this group. 45 46 47 48 366 49 50 51 367 5.2. Interpretation of the fauna from the Pachycardientuffe 52 53 368 54 55 56 57 369 Several lines of evidence indicate that the fauna of the Pachycardientuffe is allochthonous: (1) the 58 59 370 lack of fossils that were preserved in life position; (2) the lack of bivalves that are preserved with 60 61 62 63 16 64 65 371 conjoined valves; (3) frequent fragmentation of fossils; (4) the poorly sorted sediment. These 1 2 372 observations raise the question where the transported fossils originally came from. 3 4 5 6 373 In contrast to the Schlernplateau fauna, where many species occur in similar abundances, the 7 8 374 fauna from the Pachycardientuffe is clearly dominated by only two species, namely Pachycardia 9 10 375 rugosa and Pachycardia plieningeri. A comparison between the fauna of the Schlernplateau beds and 11 12 13 376 the Pachycardientuffe revealed that 45% of all species occur exclusively in the Pachycardientuffe and 14 15 377 38% of all species occur exclusively in the Schlernplateau beds. The remaining 17% of all species 16 17 378 occur in the Schlernplateau beds as well as in the Pachycardientuffe (Online Resource 1). Species that 18 19 20 379 occur in the Schlernplateau beds as well as in the Pachycardientuffe include the bivalve species 21 22 380 Myophoria kefersteini, Heminajas fissidentata, Pachycardia rugosa, Myophoriopis richthofeni and 23 24 25 381 Trigonodus rablensis, which belong to the most abundant species in the Schlernplateau beds as well 26 27 382 as in the Pachycardientuffe. All these species were shallow infaunal suspension-feeders that required 28 29 383 a soft substratum. However, there are large differences in the relative abundances of these species 30 31 32 384 between the Schlernplateau beds and the Pachycardientuffe. Myophoria kefersteini and Heminajas 33 34 385 fissidentata are the most common species in the Schlernplateau beds but only occur rarely in the 35 36 386 Pachycardientuffe. In contrast, Pachycardia rugosa is the dominant species in the Pachycardientuffe, 37 38 39 387 but it ranks third in the Schlernplateau beds, where it is much less common than Myophoria 40 41 388 kefersteini and Heminajas fissidentata. If the Schlernplateau beds were the only source area for the 42 43 389 fossils of the Pachycardientuffe, then Pachycardia rugosa should be less dominant. It is therefore 44 45 46 390 likely that the fauna of the Pachycardientuffe did not originate solely from the Schlernplateau beds. 47 48 49 391 Based on palaeogeographic reconstructions (Brandner 1991, p. 12, Fig. 6; own observations) and 50 51 392 ecological features of the fauna, we identified two additional possible source areas for the fauna of 52 53 54 393 the Pachycardientuffe. One of these additional source areas is the shallow clastic coast of a nearby 55 56 394 volcanic island and/or adjacent submarine highs, where a fauna could have flourished in 57 58 59 395 volcanoclastic sediments. Brandner (1991, p. 12, Fig. 6) indicated the presence of a structural high 60 61 396 with a volcanic island topography few kilometres southeast of the Frommerbach localities, which was 62 63 17 64 65 397 caused by the tectonic activity during the Late Ladinian and consisted of uplifted volcanoclastics of 1 2 398 the Wengen Fm. Thick shelled, shallow-burrowing bivalves such as Pachycardia are typical faunal 3 4 5 399 elements of such shallow-marine, sandy sediments (Stanley 1970). Brandner (1991, p. 12, Fig. 6) 6 7 400 indicated dipping of the palaeo-relief of the structural high towards the northwest, directly pointing 8 9 401 towards the Frommerbach localities (see also Yose 1991, p. 35). We assume that at least parts of the 10 11 12 402 fauna of the Pachycardientuffe were transported from this source area via submarine high-density 13 14 403 gravity flows, together with volcanogenic material (Figs 12a-c). A similar scenario was already 15 16 17 404 described by Fürsich and Wendt (1977), who assumed that the high number of Pachycardia indicates 18 19 405 a distinct association in a shallow-water high-energy environment, e.g. in the vicinity of volcanic 20 21 406 swells or islands, and that this Pachycardia dominated fauna together with tuffitic material and 22 23 24 407 faunal elements from neighbouring biotopes was transported into the basin. 25 26 27 408 A third possible source area is the reef environment that fringed the lagoon of the Schlernplateau 28 29 409 beds. Blaschke (1905, p. 219) reported the presence of several bivalve taxa in the Pachycardientuffe 30 31 32 410 that have previously been reported from the reefal facies of the “Schlerndolomit” sensu lato. These 33 34 411 bivalves include Chlamys (Granulochlamys) cf. tubulifera (Münster, 1841), Mysidioptera cf. 35 36 412 aviculaeformis Broili, 1903, Mysidioptera cf. incurvostriata (Gümbel, 1861), Mysidioptera cf. spinigera 37 38 39 413 Bittner, 1895, Mysidioptera acuta Broili, 1903, Mysidioptera cf. laczkói Bittner, 1901, Cassianella 40 41 414 decussata (Münster, 1841), Waagenoperna cf. planata (Broili, 1903), Pteria aff. seisiana (Broili, 1903) 42 43 415 and Badiotella sp., which were mostly epibyssate suspension-feeders that needed a hard substratum 44 45 46 416 for attachment. Furthermore, gastropods from the family Patellidae were reported from the 47 48 417 Pachycardientuffe (Blaschke 1905), which also live on hard substratum. 49 50 51 418 Fig. 13 provides a summary of our interpretation of the fauna from the Pachycardientuffe, 52 53 54 419 showing transportation from the following three sources: (1) the soft ground lagoonal fauna of the 55 56 420 Schlernplateu beds, (2) the hard ground reef fauna from the Schlerndolomite and (3) a Pachycardia 57 58 59 421 rugosa dominated fauna from the coast from a near volcanic island. 60 61 62 63 18 64 65 422 As mentioned above, the number of species found in this study (59 species) is much lower than 1 2 423 the number of species described by Broili (1903), Blaschke (1905), Waagen (1903) and Waagen 3 4 5 424 (1907; altogether more than 270 species). One explanation for the high number of species described 6 7 425 in these monographs is taxonomic oversplitting. A taxonomic revision of the fauna is beyond the 8 9 426 10 scope of this study, but an inspection of Broili’s (1903) type material in Munich confirms that his 11 12 427 taxonomic approach was more typological than usually applied today, and that e.g. the species 13 14 428 richness of the limid bivalve genus Mysidioptera (24 reported species) has been clearly 15 16 17 429 overestimated. However, also the number of reported genera is higher in the literature than in our 18 19 430 samples (100 versus 38 genera), thus oversplitting of species is not the sole reason for the difference 20 21 431 in reported diversity. An additional factor that contributed to the higher species richness in the 22 23 24 432 historical monographs is probably that these were based on a higher number of geographically 25 26 433 distant sampling localities, which likely had different source areas and thus differed considerably in 27 28 434 their taxonomic composition. Therefore, the lower species richness in our samples might by largely 29 30 31 435 due to the poorer present-day outcrop conditions, which limited our field collections to two nearby 32 33 436 localities. 34 35 36 437 37 38 39 40 41 438 6. Summarized interpretation of the benthic faunas from the Schlernplateau beds and 42 43 439 Pachycardientuffe 44 45 46 440 47 48 49 441 Our field study led to the recognition of 52 species in the Schlernplateau beds and 59 species in 50 51 442 the Pachycardientuffe. The sedimentological and palaeontological analyses outlined above allow the 52 53 54 443 detailed palaeoecological interpretation of these faunas. 55 56 57 444 The recorded species from the Schlernplateau beds lived in a lagoon, close to an oceanic inlet. The 58 59 445 fauna is unaffected from mixing by transportation, but it was probably subject to some time- 60 61 62 63 19 64 65 446 averaging. This means that some of the recorded species did not live contemporaneously at the same 1 2 447 place, thus the overall number of species is a certain overestimation of the community species 3 4 5 448 richness at a given time. However, the overall diversity of the Schlernplateau fauna represents a 6 7 449 good estimation of the taxonomic richness of shelly soft-bottom dwellers in a shallow-water setting 8 9 450 of a tropical sea at the end of the Middle Triassic. 10 11 12 13 451 In contrast to the Schlernplateau beds, the fauna of the Pachycardientuffe represents almost 14 15 452 certainly palaeocommunities from at least three different settings (lagoon, reef and sandy shallow 16 17 453 sea) that were mixed during transportation into the basin. The fact that our collection contains 18 19 20 454 significantly less species (59) than recorded in the literature (>270) is largely due to the lower 21 22 455 number of sampled localities that encompass a lower number of ecologically different source areas. 23 24 25 456 The evidence for faunal mixing in the Pachycardientuffe does not invalidate the data for studies of 26 27 28 457 biodiversity during the Middle Triassic, but it poses some constraints on their interpretation. The 29 30 458 fauna of the Pachycardientuffe does not represent a single palaeocommunity, but an integral of the 31 32 33 459 shallow marine shelly fauna that lived in different habitats at the same time. It is thus a good 34 35 460 estimator for the gamma-diversity of the reef-to-basin landscape of the tropical Tethyan ocean 36 37 461 during the late Middle Triassic. 38 39 40 41 462 42 43 44 45 463 7. Species richness of Early and Middle Triassic benthic faunas: a comparison 46 47 464 48 49 50 51 465 To what extend do Early and Middle Triassic benthic communities differ in their ecology, species 52 53 466 richness and taxonomic composition? Answering this question is crucial for identifying evolutionary 54 55 467 processes during the main phase of rediversification after the end-Permian mass extinction. We here 56 57 58 468 present a preliminary comparison between selected benthic communities of these two epochs. Data 59 60 469 used for this comparison are shown in Tab. 1. Lower and Middle Triassic macroinvertebrate faunas 61 62 63 20 64 65 470 selected for the comparison come from lithological units that represent similar shallow-marine 1 2 471 environments. Because most of the data used for this comparison are raw data from the literature, 3 4 5 472 synonyms and taxonomic oversplitting (see also chapter 5.2) may cause a certain bias towards 6 7 473 greater richness, especially in Middle Triassic faunas where modern revisions mostly lack. A 8 9 474 reevaluation of bivalve species from the Jena Fm. and the Meissner Fm. (Hautmann, unpublished 10 11 12 475 data) suggests oversplitting by nearly 50%. However, because oversplitting is likely similar in all major 13 14 476 taxa, the gastropod/bivalve ratio discussed below is probably robust, and a pronounced Anisian rise 15 16 17 477 in diversity maintains even if an overestimation of diversity by factor 2 is assumed. 18 19 20 478 The comparison between Early and Middle Triassic benthic communities shows clearly that the 21 22 479 overall diversity increased through time; however, diversity slowly increased during the Early Triassic 23 24 25 480 and rapidly during the Middle Triassic (Tab. 1; Fig. 14a). This pattern is evident in the trends of the 26 27 481 most diverse taxa in Early and Middle Triassic benthic communities, i.e., bivalves, gastropods and 28 29 482 brachiopods. Our compilation shows that bivalves were clearly the most diverse taxon in Early 30 31 32 483 Triassic benthic communities. Although bivalves were relatively diverse immediately after the end- 33 34 484 Permian mass extinction, the net increase in species richness was small until the end of the Early 35 36 485 Triassic. The major diversification of bivalves occurred during the Anisian (Hautmann 2007; Tu et al. 37 38 39 486 2016). Already the Late Bithynian–Early Illyrian Jena Fm. yields a highly diverse Middle Triassic 40 41 487 bivalve fauna, surpassed by the late Anisian fauna of the Leidapo Mb. (Tab. 1). Gastropods were on 42 43 488 average the second most diverse taxon during the Early Triassic, and their increase in species 44 45 46 489 richness during that time was comparable to that of bivalves. During the Middle Triassic, however, 47 48 490 gastropod species richness increased more rapidly than bivalve species richness. In consequence, 49 50 51 491 gastropods were the most diverse group in many Middle Triassic lithological units (Tab. 1). The 52 53 492 change from the taxonomic dominance of bivalves in Early Triassic benthic communities to the 54 55 493 taxonomic dominance of gastropods during the Middle Triassic can also be seen in the 56 57 58 494 gastropod/bivalve-ratio that was variable but with an increasing trend through time (Fig. 14b). 59 60 495 Brachiopods were the third most diverse group in almost all Early and Middle Triassic benthic 61 62 63 21 64 65 496 communities. Their species richness was relatively low during the Early Triassic, and in most 1 2 497 Olenekian lithological units used for this comparison they are even not reported. However, 3 4 5 498 brachiopods also became more diverse during the Middle Triassic, although their increase in species 6 7 499 richness during the Middle Triassic is much slower compared to bivalves and gastropods. 8 9 10 500 The comparison between Early and Middle Triassic benthic communities confirms that corals 11 12 13 501 were absent in the entire Early Triassic (e.g., Flügel 2002; Weidlich et al. 2003; Stanley 2003; Payne et 14 15 502 al. 2006; Brayard et al. 2011). Sponges occurred only in two Lower Triassic formations used for this 16 17 503 comparison, but sponges, in association with microbialites, are known from the Lower Triassic 18 19 20 504 (Induan) of Armenia (Friesenbichler et al. 2018), Lower and Middle Triassic carbonate successions of 21 22 505 the Western Tethys domain (the Alps, Carpathians and the Germanic Basin; Szulc 2007) as well as 23 24 25 506 Lower Triassic strata from China (Ezaki et al. 2008; Bagherpour et al. 2017) and the Western US 26 27 507 (Brayard et al. 2011; Hofmann et al. 2014). However, the number of these sponge species is not 28 29 508 known. Middle Triassic sponges are not only reported in the lithological units shown in Tab. 1 but 30 31 32 509 also in the Illyrian strata of the Wetterstein Fm. (Velledits et al. 2011). The fossil record of 33 34 510 echinoderms in the Early to Middle Triassic is heterogeneous. Several Palaeozoic clades of 35 36 511 echinoderms survived the end-Permian mass extinction and reappeared as Lazarus taxa in the 37 38 39 512 Middle Triassic, without giving rise to a new diversification (Thuy et al. 2017). Crinoid diversity was 40 41 513 very low during the Early Triassic (Hagdorn 2011), although cladistics analyses suggest that initial 42 43 514 diversification took place in the Induan (Twitchett and Oji 2005) or already in the Late Permian (Oji 44 45 46 515 and Twitchett (2015); however, well before the documented species richness increase during the 47 48 516 Middle and early Late Triassic (Hagdorn 2011). Only two echinoid genera are reported from Lower 49 50 51 517 Triassic strata (Twitchett and Oji 2005) and very low levels of diversity were apparently maintained 52 53 518 for at least the duration of the Early and Middle Triassic. Nevertheless, numerous undescribed 54 55 519 echinoid spines and other remains have been noted in Early Triassic fossil assemblages (e.g., from the 56 57 58 520 Griesbachian of Oman; Krystyn et al. 2003) and these undescribed records may hint at a hidden 59 60 521 diversity of echinoids within the Early Triassic (see also Thompson et al. 2015; Thuy et al. 2017). Fossil 61 62 63 22 64 65 522 remains or trace fossils attributed to Lower Triassic asteroids have only been recorded from 1 2 523 Olenekian strata of the western USA (Schubert and Bottjer 1995; Villier et al. 2018). Definite reports 3 4 5 524 from Lower Triassic holothuroids only exist from Western Nepal (Kozur and Mostler 1973; Mostler 6 7 525 1977), whereas a total number of 84 holothuroid species was reported from several Middle Triassic 8 9 526 strata (Reich 2013). Fossil remains from ophiuroids are quite abundant in Lower Triassic strata 10 11 12 527 worldwide but so far only six species have been described (Twitchett and Oji 2005). Early Triassic 13 14 528 bryozoans were reported from a couple of Lower Triassic strata (Nakrem and Mørk 1991; Wignall and 15 16 17 529 Twitchett 2002; Baud et al. 2008) but only five Early Triassic bryozoan genera are reported (Powers 18 19 530 and Pachut 2007) and only six species have been described formally (Baud et al. 2008); however, 20 21 531 bryozoan diversity increased during the Middle and Late Triassic, with eight reported genera (Powers 22 23 24 532 and Pachut 2007) a total number of more than 40 species (see reviews in Schäfer and Grant-Mackie 25 26 533 1994; Schäfer et al. 2003). Microconchids were important constituents of marine ecosystems 27 28 534 immediately after the Permian/Triassic mass extinction (He et al. 2012) and were also commonly 29 30 31 535 present through the entire Early Triassic worldwide (Fraiser 2011; Zatoń et al. 2013b; Yang et al. 32 33 536 2015). Although microconchids can be very abundant locally, they are lowly diverse and only 34 35 36 537 represented by three genera (Zatoń et al. 2013b). 37 38 39 538 The Middle Triassic increase in species richness might be related to more stable environmental 40 41 539 conditions, as reflected by the constancy of the Middle Triassic carbon isotope curve (Payne et al. 42 43 540 2004), or alternatively represent the effect of competition-driven niche partitioning after 44 45 46 541 rediversification exceeded a threshold (Hautmann 2014; Hautmann et al. 2015). The change in the 47 48 542 gastropod/bivalve-ratio is more difficult to interpret, but it is notable that occurrences of high 49 50 51 543 gastropod species richness relative to bivalves are almost exclusively linked to carbonate platform 52 53 544 environments (Fig. 14b). This might represent a taphonomic bias due to differences in skeletal 54 55 545 mineralogy between gastropods and bivalves, but we note that aragonite-shelled bivalves are 56 57 58 546 commonly preserved in the Early Triassic, which makes this explanation unlikely. Alternatively, we 59 60 547 hypothesise that the increase in gastropod species richness is a primary signal linked to the 61 62 63 23 64 65 548 resurgence of large carbonate platform and reefs during the Middle Triassic (Gaetani et al. 1981; 1 2 549 Senowbari et al. 1993), although details of the ecological causes for this facies-related increase in 3 4 5 550 gastropod species richness requires further investigation. 6 7 8 551 9 10 11 12 552 8. Conclusion 13 14 15 553 16 17 18 554 Early and Middle Triassic shallow-marine benthic communities differed considerably in terms of 19 20 555 species richness and taxonomic composition. Whereas Early Triassic formations from shallow-marine 21 22 23 556 settings contain generally less than 50 macroinvertebrate species, regional diversities of around 300 24 25 557 species have been described from various Anisian-Ladinian shallow-marine settings worldwide (Tab. 26 27 28 558 1; Fig. 14), including the herein studied fauna from the Pachycardientuffe. For more narrowly defined 29 30 559 habitats, such as that of the open lagoon represented by the Schlernplateau fauna (85 species), the 31 32 560 increase in diversity was likewise remarkable. The Middle Triassic increase in diversity at the 33 34 35 561 local/regional level analysed herein correlates with the most pronounced increase in bivalve diversity 36 37 562 during the entire Triassic in a global dataset (Hautmann 2007, fig. 8) and therefore represents the 38 39 563 main phase of recovery after the end-Permian mass extinction. This main phase of recovery in 40 41 42 564 benthic marine communities was associated with a weakening of the dominance of bivalves in 43 44 565 comparison to the Early Triassic, which reflects the proliferation of other invertebrate taxa, 45 46 566 particularly gastropods. We hypothesise that the resurgence of carbonate platforms during the 47 48 49 567 Middle Triassic played an important role in the shift of taxonomic compositions, and that the 50 51 568 increase of environmental diversity associated with newly arising carbonate platforms contributed at 52 53 54 569 least partly to the accelerated pace of recovery during the Middle Triassic. 55 56 57 58 59 60 61 62 63 24 64 65 570 References 1 2 571 3 4 5 6 572 Aberhan, M. 1994. Guild-Structure and evolution of Mesozoic Benthic shelf Communities. Palaios 9: 7 8 573 516–545. 9 10 11 574 Alroy, J., M. Aberhan, D.J. Bottjer, M. Foote, F.T. Fürsich, P.J. Harries, A.J.W. Hendy, S.M. Holland, L.C. 12 13 14 575 Ivany, W. Kiessling, M.A. Kosnik, C.R. Marshall, A.J. McGowan, A.I. Miller, T.D. Olszewski, M.E. 15 16 576 Patzkowsky, S.E. Peters, L. Villier, P.J. Wagner, N. Bonuso, P.S. Borkow, B. Brenneis, M.E. Clapham, 17 18 577 L.M. Fall, C.A. Ferguson, V.L. Hanson, A.Z. Krug, K.M. Layou, E.H. Leckey, S. Nürnberg, C.M. 19 20 21 578 Powers, J.A. Sessa, C. Simpson, A. Tomašových, and C.C. Visaggi. 2008. Phanerozoic trends in the 22 23 579 global diversity of marine invertebrates. Science 321: 97–100. 24 25 26 580 Autonome Provinz Bozen (ed.). 2007. Geologische Karte der Westlichen Dolomiten. 27 28 29 30 581 Bagherpour, B., H. Bucher, A. Baud, M. Brosse, T. Vennemann, R. Martin, and K. Guodun. 2017. 31 32 582 Onset, development, and cessation of basal Early Triassic microbialites (BETM) in the Nanpanjiang 33 34 583 pull-apart Basin, South China Block. Gondwana Research 44: 178–204. 35 36 37 38 584 Balini, M., S.G. Lucas, J.F. Jenks, and J.A. Spielmann. 2010. Triassic ammonoid biostratigraphy: an 39 40 585 overview. In The Triassic Timescale, ed. S.G. Lucas, 221–262. Geological Society of London, Special 41 42 586 Publications 334. 43 44 45 46 587 Batten, R.L., and W.L. Stokes. 1986. Early Triassic gastropods from the Sinbad Member of the 47 48 588 Moenkopi Formation, San Rafael Swell, Utah. American Museum Novitates 2864: 1–33. 49 50 51 589 Baud, A., H.A. Nakrem, B. Beauchamp, T.W. Beatty, A.F. Embry, and C.M. Henderson. 2008. Lower 52 53 54 590 Triassic bryozoan beds from Ellesmere Island, High Arctic, Canada. Polar Research 27: 428–440. 55 56 57 591 Bittner, A. 1890. Brachiopoden der alpinen Trias. Abhandlungen der k. k. geologischen Reichsanstalt 58 59 592 14: 1–325. 60 61 62 63 25 64 65 593 Bittner, A. 1892. Brachiopoden der alpinen Trias - Nachtrag. Abhandlungen der k. k. geologischen 1 2 594 Reichsanstalt 17: 1–55. 3 4 5 6 595 Bittner, A. 1895. Lamellibranchiaten der alpinen Trias. I Theil: Revision der Lamellibranchiaten von 7 8 596 Sct. Cassian. Abhandlungen der k. k. Geologischen Reichsanstalt 18: 1–236. 9 10 11 597 Bittner, A. 1900. Ueber die triadische Lamellibranchiaten-Gattung Mysidioptera Sal. und deren 12 13 14 598 Beziehungen zu palaeozoischen Gattungen. Jahrbuch der k. k. Geologischen Reichsanstalt 50: 59– 15 16 599 66. 17 18 19 600 Bittner, A. 1901. Lamellibranchiaten aus der Trias des Bakonyer Waldes. Resultate der 20 21 22 601 wissenschaftlichen Erforschung des Balatonsees, Anhang: Palaeontologie der Umgebung des 23 24 602 Balatonsees II, Band 3: 1–107. 25 26 27 603 Blaschke, F. 1905. Die Gastropodenfauna der Pachycardientuffe der Seiseralpe in Südtirol nebst 28 29 30 604 einem Nachtrag zur Gastropodenfauna der roten Raibler Schichten auf dem Schlernplateau. 31 32 605 Beiträge zur Paläontologie und Geologie Österreich-Ungarns und des Orients 17: 161–221. 33 34 35 606 Böhm, J. 1895. Die Gastropoden des Marmolatakalkes. Palaeontographica 42: 211–308. 36 37 38 607 Bosellini, A. 1998. Geologie der Dolomiten. Bozen: Verlagsanstalt Athesia. 39 40 41 42 608 Bottoli, G., and S. Trombetta. 1998. Analisi di facies ed ambiente deposizionale del Conglomerato 43 44 609 della Marmolada: un sistema torbiditico dominato da flussi ad alta densità (gruppo Col Rossi-Porta 45 46 610 Vescovo, Dolomti occidentali). Memorie Società Geologica Italiana 53: 341 357. 47 – 48 49 50 611 Brack, P., and H. Rieber. 1993. Toward a better definition of the Anisian/Ladinian boundary: New 51 52 612 biostratigraphic data and correlations of boundary sections from the Southern Alps. Eclogae 53 54 613 Geologicae Helvetiae 86: 415 527. 55 – 56 57 58 59 60 61 62 63 26 64 65 614 Brack, P., R. Mundil, F. Oberli, M. Meier, and H. Rieber. 1996. Biostratigraphic and radiometric age 1 2 615 data question the Milankovitch characteristics of the Latemar cycles (Southern Alps, Italy). 3 4 5 616 Geology 24: 371–375. 6 7 8 617 Brandner, R. 1991. Geological Setting and Stratigraphy of the Schlern-Rosengarten Buildup and Seiser 9 10 618 Alm Basin. In The Northern Margin of the Schlern/Sciliar-Rosengarten/Catinaccio Platform, eds. R. 11 12 13 619 Brandner, E. Flügel, R. Koch, and L.A. Yose, 4–16. Dolomieu Conference on Carbonate Platforms 14 15 620 and Dolomitization, Guidebook Excursion A, Ortisei (Italy). 16 17 18 621 Brandner, R., A. Gruber, and L. Keim. 2007. Geologie der westlichen Dolomiten: Von der Geburt der 19 20 21 622 Neotethys im Perm zu Karbonatplattformen, Becken und Vulkaniten der Trias. Geo.Alp 4: 95–121. 22 23 24 623 Brandner, R., A. Gruber, C. Morelli, and V. Mair. 2016. Pulses of Neotethys-Rifting in the 25 26 624 Permomesozoic of the Dolomites. Geo.Alp 13: 7–70. 27 28 29 30 625 Brayard, A., E. Vennin, N. Olivier, K G. Bylund, J. Jenks, D.A. Stephen, H. Bucher, R. Hofmann, N. 31 32 626 Goudemand, and G. Escarguel. 2011. Transient metazoan reefs in the aftermath of the end- 33 34 627 Permian mass extinction. Nature Geoscience 4: 693–697. 35 36 37 38 628 Brayard, A., G. Escarguel, H. Bucher, C. Monnet, T. Brühwiler, N. Goudemand, T. Galfetti, and J. Guex. 39 40 629 2009. Good genes and good luck: ammonoid diversity and the end-Permian mass extinction. 41 42 630 Science 325: 1118–1121. 43 44 45 46 631 Brayard, A., L.J. Krumenacker, J.P. Botting, J.F. Jenks, K.G. Bylund, E. Fara, E. Vennin, N. Olivier, N. 47 48 632 Goudemand, T. Saucède, S. Charbonnier, C. Romano, L. Doguzhaeva, B. Thuy, M. Hautmann, D.A. 49 50 633 Stephen, C. Thomazo, G. and Excarguel. 2017. Unexpected Early Triassic marine ecosystem and 51 52 53 634 the rise of the Modern fauna. Science Advances 3: e1602159. 54 55 56 635 Broglio Loriga, C., C. Neri, and R. Posenato. 1986. The Early Macrofaunas of the Werfen Formation 57 58 636 and the Permian-Triassic Boundary in the Dolomites (Southern Alps, Italy). Acta Geologica 62: 3– 59 60 61 637 18. 62 63 27 64 65 638 Broglio Loriga, C., F. Góczán, J. Haas, K. Lenner, C. Neri, A. Oravecz-Scheffer, R. Posenato, I. Szabó, 1 2 639 and Á. Tóth-Makk. 1990. The Lower Triassic sequences of the Dolomites (Italy) and Transdanubian 3 4 5 640 Mid-Mountains (Hungary) and their correlation. Memorie di Scienze Geologiche 42: 41–103. 6 7 8 641 Broili, F. 1903. Die Fauna der Pachycardientuffe der Seiser Alp (Mit Ausschluss der Gastropoden und 9 10 642 Cephalopoden). Palaeontographica 50: 145–227. 11 12 13 14 643 Budai, T. 2008. Geological excursions in the Alpine and Germanic Triassic facies areas of Hungary. 15 16 644 Triassic Workshop 2008, Hungary: 1–50. 17 18 19 645 Busse, E. 1972. Fazies und Fauna des Oberen Muschelkalkes von Willebadessen. Philippia 1: 110–126. 20 21 22 646 Eichwald, E. 1851. Naturhistorische Bemerkungen als Beitrag zur vergleichenden Geognosie, auf 23 24 25 647 einer Reise durch die Eifel, Tyrol, Italien, Sizilien und Algier. II. Geognostischer Ausflug nach Tyrol. 26 27 648 Nouveaux mémoires de la Société impériale des naturalistes de Moscou 9: 75–204. 28 29 30 649 Ezaki Y., J. Liu, T. Nagano and N. Adachi. 2008. Geobiological aspects of the earliest Triassic 31 32 33 650 microbialites along the southern periphery of the tropical Yangtze Platform: initiation and 34 35 651 cessation of a microbial regime. Palaios 23: 356–369. 36 37 38 652 Flügel, E. 2002. Triassic reef patterns. In Phanerozoic Reef Patterns, eds. W. Kiessling, E. Flügel, and J. 39 40 41 653 Golonka, 391–463. SEPM Special Publication 72, Society for Sedimentary Geology. 42 43 44 654 Foster, W.J., and K. Sebe. 2017. Recovery and diversification of marine communities following the 45 46 655 late Permian mass extinction event in the western Palaeotethys. Global and Planetary Change 47 48 49 656 155: 165–177. 50 51 52 657 Foster, W.J., S. Danise, A. Sedlacek, G.D. Price, K. Hips, and R.J. Twitchett. 2015. Environmental 53 54 658 control on the post-Permian recovery of benthic, tropical marine ecosystems in western 55 56 57 659 Palaeotethys (Aggtelek Karst, Hungary). Palaeogeography, Palaeoclimatology, Palaeoecology 440: 58 59 660 374–394. 60 61 62 63 28 64 65 661 Foster, W.J., S. Danise, G.D. Price, and R.J. Twitchett. 2017. Subsequent biotic crises delayed marine 1 2 662 recovery following the late Permian mass extinction event in northern Italy. PLoS One 12: 3 4 5 663 e0172321. 6 7 8 664 Fraiser, M.L. 2011. Paleoecology of second tiers from western Pangean tropical marine environments 9 10 665 during the aftermath of the end-Permian mass extinction. Palaeogeography, Palaeoclimatology, 11 12 13 666 Palaeoecology 308: 181–189. 14 15 16 667 Friesenbichler, E., S. Richoz, A. Baud, L. Krystyn, L. Sahakyan, S. Vardanyan, J. Peckmann, J. Reitner, 17 18 668 and K. Heindel. 2018. Sponge-microbial build-ups from the lowermost Triassic Chanakhchi section 19 20 21 669 in southern Armenia: Microfacies and stable carbon isotopes. Palaeogeography, 22 23 670 Palaeoclimatology, Palaeoecology 490: 653–672. 24 25 26 671 Frizzell, D.L., and H. Exline. 1956. Monograph of Fossil Holothurian Sclerites. Bulletin of School of 27 28 29 672 Mines and Metallurgy (Technical Series) 89: 1–204. 30 31 32 673 Fürsich, F.T. 1994. Palaeoecology and evolution of Mesozoic salinity-controlled benthic 33 34 674 macroinvertebrate associations. Lethaia 26: 327–346. 35 36 37 38 675 Fürsich, F.T., and J. Wendt. 1977. Biostratinomy and palaeoecology of the Cassian Formation 39 40 676 (Triassic) of the Southern Alps. Palaeogeography, Palaeoclimatology, Palaeoecology 22: 257–323. 41 42 43 677 Fürsich, F.T., and M. Aberhan. 1990. Significance of time‐averaging for palaeocommunity analysis. 44 45 46 678 Lethaia 23: 143–152. 47 48 49 679 Fürsich, F.T., S. Freytag, J. Röhl, and A. Schmid. 1995. Palaeoecology of benthic associations in 50 51 680 salinity-controlled marginal marine environments: Examples from the Lower Bathonian (Jurassic) 52 53 54 681 of the Causses (southern France). Palaeogeography, Palaeoclimatology, Palaeoecology 113: 135– 55 56 682 172. 57 58 59 60 61 62 63 29 64 65 683 Gaetani, M., E. Fois, F. Jadoul, and A. Nicora. 1981. Nature and evolution of Middle Triassic carbonate 1 2 684 buildups in the Dolomites (Italy). Marine Geology 44: 25–57. 3 4 5 6 685 Goldfuss, G.A. 1833-1841. Petrefacta Germaniae. Teil 2. 1833: pp. 1–68, pls. 72–96; 1835: pp. 69– 7 8 686 140, pls. 97–121; 1837: pp. 141–224, pls. 122–146; 1841: pp. 225–312, pls. 147–165; Düsseldorf 9 10 687 (Arnz & Co). 11 12 13 14 688 Gredler, V.M. 1862. Vierzehn Tage in Bad Ratzes: eine naturgeschichtliche Lokalskizze mit näherer 15 16 689 Berücksichtigung der Fauna. Programm des Kaiserlich-Königlichen Gymnasiums zu Bozen, 17 18 690 veröffentlicht am Schlusse des Schuljahres 13 (1862/63): 1–41. 19 20 21 22 691 Gümbel, C.W. 1861. Geognostische Beschreibung des bayerischen Alpengebirges und seines 23 24 692 Vorlandes. Gotha: Verlag von Justus Perthes. 25 26 27 693 Häberle, D. 1908. Paläontologische Untersuchung triadischer Gastropoden aus dem Gebiet von 28 29 30 694 Predazzo. Verhandlungen des Naturhistorisch-Medizinischen Vereins zu Heidelberg 9: 1–630. 31 32 33 695 Hagdorn, H. 2011. Triassic: the crucial period of post-Palaeozoic crinoid diversification. Swiss Journal 34 35 696 of Palaeontology 130: 91–122. 36 37 38 697 Hagdorn, H., and E. Nitsch. 2009. 6th International Triassic Field Workshop (Pan-European Correlation 39 40 41 698 of the Triassic). Triassic of Southwest Germany. Freiburg: Landesamt für Geologie, Rohstoffe und 42 43 699 Bergbau. 44 45 46 700 Hagdorn, H., and R. Mundlos. 1982. Autochthonschille im Oberen Muschelkalk (Mitteltrias) 47 48 49 701 Südwestdeutschlands. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 162: 332– 50 51 702 351. 52 53 54 703 Hagdorn, H., and W. Ockert. 1993. Encrinus liliiformis im Trochitenkalk Süddeutschlands. In: 55 56 57 704 Muschelkalk. Sonderbände der Gesellschaft für Naturkunde in Württemberg e.V. 2, 245–260. 58 59 705 Stuttgart, Korb: Goldschneck-Verlag Werner K. Weidert. 60 61 62 63 30 64 65 706 Hammer, Ø., D.A.T. Harper, and P.D. Ryan. 2001. PAST: Paleontological Statistics Software Package 1 2 707 for Education and Data Analysis. Palaeontologia Electronica 4: 1–9. 3 4 5 6 708 Hauer, F. von. 1857. Ein Beitrag zur Kenntniss der Fauna der Raibler Schichten. Sitzungsberichte der 7 8 709 Kaiserlichen Akademie der Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klasse 9 10 710 24: 1–32. 11 12 13 14 711 Hausmann, I.M., A. and Nützel. 2015. Diversity and palaeoecology of a highly diverse Late Triassic 15 16 712 marine biota from the Cassian Formation of north Italy. Lethaia 48: 235–255. 17 18 19 713 Hautmann, M. 2007. Steuerungsfaktoren globaler Biodiversifizierungsmuster am Beispiel 20 21 22 714 frühmesozoischer Muscheln. Beringeria 37: 61–74. 23 24 25 715 Hautmann, M. 2014. Diversification and diversity partitioning. Paleobiology 40: 162–176. 26 27 28 716 Hautmann, M., A.B. Smith, A.J. McGowan, and H. Bucher. 2013. Bivalves from the Olenekian (Early 29 30 717 Triassic of south-western Utah: systematics and evolutionary significance. Journal of Systematic 31 32 33 718 Palaeontology 11: 263–293. 34 35 36 719 Hautmann, M., and H. Hagdorn. 2013. Oysters and oyster-like bivalves from the Middle Triassic 37 38 720 Muschelkalk of the Germanic Basin. Paläontologische Zeitschrift 87: 19 32. 39 – 40 41 42 721 Hautmann, M., B. Bagherpour, M. Brosse, Å. Frisk, R. Hofmann, A. Baud, A. Nützel, N. Goudemand, 43 44 722 and H. Bucher. 2015. Competition in slow motion: The unusual case of benthic marine 45 46 723 communities in the wake of the end-Permian mass extinction. Palaeontology 58: 871 901. 47 – 48 49 50 724 Hautmann, M., H. Bucher, T. Brühwiler, N. Goudemand, A. Kaim, and A. Nützel. 2011. An unusually 51 52 725 diverse mollusc fauna from the earliest Triassic of South China and its implications for benthic 53 54 726 recovery after the end-Permian biotic crisis. Geobios 44: 71 85. 55 – 56 57 58 727 He, L., Y.B. Wang, A. Woods, G.S. Li, H. Yang, and W. Liao. 2012. Calcareous tubeworms as disaster 59 60 728 forms after the end-Permian mass extinction in South China. Palaios 27: 878–886. 61 62 63 31 64 65 729 Hofmann, R., M. Hautmann, A. Brayard, A. Nützel, K.G. Bylund, J.F. Jenks, E. Vennin, N. Olivier, and H. 1 2 730 Bucher. 2014. Recovery of benthic marine communities from the end‐Permian mass extinction at 3 4 5 731 the low latitudes of eastern Panthalassa. Palaeontology 57: 547–589. 6 7 8 732 Hofmann, R., M. Hautmann, and H. Bucher. 2013a. A new paleoecological look at the Dinwoody 9 10 733 Formation (Lower Triassic, Western USA): Intrinsic versus extrinsic controls on ecosystem 11 12 13 734 recovery after the end-Permian mass extinction. Journal of Paleontology 87: 854–880. 14 15 16 735 Hofmann, R., M. Hautmann, and H. Bucher. 2015. Recovery dynamics of benthic marine communities 17 18 736 from the Lower Triassic Werfen Formation, northern Italy. Lethaia 48: 474–496. 19 20 21 22 737 Hofmann, R., M. Hautmann, M. Wasmer, and H. Bucher. 2013b. Palaeoecology of the Spathian Virgin 23 24 738 Formation (Utah, USA) and its implications for the Early Triassic recovery. Acta Palaeontologica 25 26 739 Polonica 58: 149–173. 27 28 29 30 740 Keim, L. 2008. Geologie im Gebiet Schlern-Seiser Alm: vom Tethysmeer zum Gebirge. Gredleriana 8: 31 32 741 25–46. 33 34 35 742 Kittl, E. 1891. Die Gastropoden der Schichten von St. Cassian der südalpinen Trias. I. Theil. Annalen 36 37 38 743 des Naturhistorischen Museums in Wien 6: 166–262. 39 40 41 744 Kittl, E. 1892. Die Gastropoden der Schichten von St. Cassian der südalpinen Trias. II. Theil. Annalen 42 43 745 des Naturhistorischen Museums in Wien 7: 35–97. 44 45 46 746 Kittl, E. 1894. Die Gastropoden der Schichten von St. Cassian der südalpinen Trias. III. Theil. Annalen 47 48 49 747 des Naturhistorischen Museums in Wien 9: 143–277. 50 51 52 748 Klipstein, A. von 1843-1845. Mittheilungen aus dem Gebiete der Geologie und Palaeontologie. 1843: 53 54 749 pp. 1 144, pls. 1 9; 1844: pp. 145 240, pls. 10 15; 1845: pp. 241 311, pls. 16 20; Gießen 55 – – – – – – 56 57 750 (Heyer). 3. Abt. (1883): 1–101, 3 Tafeln; Gießen (Ricker). 58 59 60 61 62 63 32 64 65 751 Komatsu, T., J.H. Chen, M.Z. Cao, F. Stiller, and H. Naruse. 2004. Middle Triassic (Anisian) diversified 1 2 752 bivalves: depositional environments and bivalve assemblages in the Leidapo Member of the 3 4 5 753 Qingyan Formation, southern China. Palaeogeography, Palaeoclimatology, Palaeoecology 208: 6 7 754 207–223. 8 9 10 755 Kozur, H., and H. Mostler. 1973. Beiträge zur Mikrofauna permotriadischer Schichtfolgen Teil 1: 11 12 13 756 Conodonten aus der Tibetzone des Niederen Himalaya (Dolpogebiet, Westnepal). Geologische 14 15 757 und Paläontologische Mitteilungen Innsbruck 3: 1–23. 16 17 18 758 Krystyn, L., A. Baud, S. Richoz, and R.J. Twitchett. 2003. A unique Permian-Triassic boundary section 19 20 21 759 from Oman. Palaeogeography, Palaeoclimatology, Palaeoecology 191: 329–344. 22 23 24 760 Laube, G.C. 1865a. Die Fauna der Schichten von St. Cassian: Ein Beitrag zur Paläontologie der alpinen 25 26 761 Trias, I. Abtheilung, Spongitarien, Corallen, Echiniden und Crinoiden. Wien: Kaiserlich-königliche 27 28 29 762 Hof- und Staatsdruckerei. 30 31 32 763 Laube, G.C. 1865b. Die Fauna der Schichten von St. Cassian: Ein Beitrag zur Paläontologie der alpinen 33 34 764 Trias, II. Abtheilung, Brachiopoden und Bivalven. Wien: Kaiserlich-königliche Hof- und 35 36 37 765 Staatsdruckerei. 38 39 40 766 Laube, G.C. 1868. Die Fauna der Schichten von St. Cassian: Ein Beitrag zur Paläontologie der alpinen 41 42 767 Trias, III. Abtheilung, Gastropoden – I. Hälfte. Wien: Kaiserlich-königliche Hof- und 43 44 45 768 Staatsdruckerei. 46 47 48 769 Laube, G.C. 1869a. Die Fauna der Schichten von St. Cassian: Ein Beitrag zur Paläontologie der alpinen 49 50 770 Trias, IV. Abtheilung, Gastropoden – II. Hälfte. Wien: Kaiserlich-königliche Hof- und 51 52 53 771 Staatsdruckerei. 54 55 56 772 Laube, G.C. 1869b. Die Fauna der Schichten von St. Cassian: Ein Beitrag zur Paläontologie der alpinen 57 58 773 Trias, V. Abtheilung, Cephalopoden. – Schluss. Wien: Kaiserlich-königliche Hof- und 59 60 61 774 Staatsdruckerei. 62 63 33 64 65 775 Leonardi, P., and F. Fiscon. 1947. Nuove forme di gasteropodi della fauna cassiana di Cortina 1 2 776 d’Ampezzo. Memorie del Museo di Storia Naturale della Venezia Tridentina 8: 1–16. 3 4 5 6 777 Leonardi, P., and F. Fiscon. 1959. La fauna cassiana di Cortina d’Ampezzo. Parte III: Gasteropodi. 7 8 778 Memorie degli Istituti di Geologia e Mineralogia dell’Università di Padova 21: 1–140. 9 10 11 779 Manfrin, S., P. Mietto, and N. Preto. 2005. Ammonoid biostratigraphy of the Middle Triassic Latemar 12 13 14 780 platform (Dolomites, Italy) and its correlation with Nevada and Canada. Geobios 38: 477–504. 15 16 17 781 McGowan, A.J., A.B. Smith, and P.D. Taylor. 2009. Faunal diversity, heterogeneity and body size in the 18 19 782 Early Triassic: testing post-extinction paradigms in the Virgin Limestone of Utah, USA. Australian 20 21 22 783 Journal of Earth Sciences 56: 859–872. 23 24 25 784 Morisita, M. 1959. Measuring interspecific association and similarity between communities. 26 27 785 Memoires of the faculty of Science Kyushu University series E (Biology) 3: 65–80. 28 29 30 786 Mostler, H. 1977. Zur Palökologie triadischer Holothurien (Echinodermata). Berichte des 31 32 33 787 naturwissenschaftlich-medizinischen Vereins in Innsbruck 64: 13–40. 34 35 36 788 Münster, G. 1841. Beiträge zur Geognosie und Petrefacten-Kunde des südöstlichen Tirols vorzüglich 37 38 789 der Schichten von St. Cassian. II. Beschreibung und Abbildung der in den Kalkmergelschichten von 39 40 41 790 St. Cassian gefunden Versteinerungen. Beiträge zur Petrefacten-Kunde 4: 25–152. 42 43 44 791 Nakrem, H.A., and A. Mørk. 1991. New Early Triassic Bryozoa (Trepostomata) from Spitsbergen, with 45 46 792 some remarks on the stratigraphy of the investigated horizons. Geological Magazine 128: 129 47 – 48 49 793 140. 50 51 52 794 Neri, C., and R. Posenato. 1985. New biostratigraphical data on uppermost Werfen Formation of 53 54 795 western Dolomites (Trento, Italy). Geologisch-Paläontologische Mitteilungen Innsbruck 14: 83 55 – 56 57 796 107. 58 59 60 61 62 63 34 64 65 797 Neyman, A.A. 1967. Limits to the application of the trophic group concept in benthic studies. 1 2 798 Oceanology 7: 149–155. 3 4 5 6 799 Nützel, A., and A. Kaim. 2014. Diversity, palaeoecology and systematics of a marine fossil assemblage 7 8 800 from the Late Triassic Cassian Formation at Settsass Scharte, N Italy. Paläontologische Zeitschrift 9 10 801 88: 405–431. 11 12 13 14 802 Nützel, A., and C. Schulbert. 2005. Facies of two important Early Triassic gastropod lagerstätten: 15 16 803 implications for diversity patterns in the aftermath of the end-Permian mass extinction. Facies 51: 17 18 804 480–500. 19 20 21 22 805 Ockert, W. 1993. Die Zwergfaunaschichten (Unterer Hauptmuschelkalk, Trochitenkalk, mo1) im 23 24 806 nordöstlichen Baden-Württemberg. In: Muschelkalk. Sonderbände der Gesellschaft für 25 26 807 Naturkunde in Württemberg e.V. 2, 117–215. Stuttgart, Korb: Goldschneck-Verlag Werner K. 27 28 29 808 Weidert. 30 31 32 809 Oji, T., and J. Twitchett. 2015. The oldest Post-Palaeozoic Crinoid and Permian-Triassic Origins of the 33 34 810 Articulata (Echinodermata). Zoological Science 32: 211–215. 35 36 37 38 811 Payne, J. 2005. Evolutionary dynamics of gastropod size across the end-Permian extinction and 39 40 812 through the Triassic recovery interval. Paleobiology 31: 269–290. 41 42 43 813 Payne, J.L., and M.E. Clapham. 2012. End-Permian mass extinction in the oceans: an ancient analog 44 45 46 814 for the twenty-first century? Annual Review of Earth and Planetary Sciences 40: 89–111. 47 48 49 815 Payne, J.L., D.J. Lehrmann, J. Wei, M.J. Orchard, D.P. Schrag, and A.H. Knoll. 2004. Large 50 51 816 perturbations of the carbon cycle during recovery from the end-Permian extinction. Science 305: 52 53 54 817 506–509. 55 56 57 58 59 60 61 62 63 35 64 65 818 Payne, J.L., D.J. Lehrmann, S. Christensen, J. Wei, and A.H. Knoll. 2006. Environmental and biological 1 2 819 controls on the initiation and growth of a Middle Triassic (Anisian) reef complex on the Great Bank 3 4 5 820 of Guizhou, Guizhou Province, China. Palaios 21: 325–343. 6 7 8 821 Pia, J. 1937. Stratigraphie und Tektonik der Pragser Dolomiten in Südtirol. Wien: Eigenverlag. 9 10 11 822 Powers, C., and J.F. Pachut. 2008. Diversity and distribution of Triassic bryozoans in the aftermath of 12 13 14 823 the end-Permian mass extinction. Journal of Paleontology 82: 362–371. 15 16 17 824 Raup, D.M. 1979. Size of the Permo-Triassic bottleneck and its evolutionary implications. Science 206: 18 19 825 217–218. 20 21 22 826 Reich, M. 2013. How many species of fossil holothurians are there? In Echinoderms in a Changing 23 24 25 827 World, ed. C. Johnson, 23–51. Proceedings of the 13th International Echinoderm Conference, 26 27 828 University of Tasmania, Hobart Tasmania, Australia. 28 29 30 829 Richthofen, F. von. 1860. Geognostische Beschreibung der Umgegend von Predazzo, Sanct Cassian 31 32 33 830 und der Seiser Alpe in Süd-Tyrol. Gotha: Perthes. 34 35 36 831 Salomon, W. 1895. Geologische und palaeontologische Studien über die Marmolata. 37 38 832 Palaeontographica 42: 1 210. 39 – 40 41 42 833 Schäfer, P., and J. Grant-Mackie. 1994. Triassic Bryozoa from the Murihiku and Torlesse Supergroups, 43 44 834 New Zealand. Memoirs of the Association of Australasian Palaeontologists 16: 1–52. 45 46 47 835 Schäfer, P., R.J. Cuffey, and A.R. Young. 2003. New trepostome Bryozoa from the Early Triassic 48 49 50 836 (Smithian/Spathian) of Nevada. Paläontologische Zeitschrift 77: 323–340. 51 52 53 837 Schmidt, M. 1928. Die Lebewelt unserer Trias. Öhringen: Hohenlohesche Buchhandlung F. Rau. 54 55 56 838 Schmidt, M. 1938. Die Lebewelt unserer Trias - Nachtrag. Öhringen: Hohenlohesche Buchhandlung F. 57 58 59 839 Rau. 60 61 62 63 36 64 65 840 Schubert, J.K., and D.J. Bottjer. 1995. Aftermath of the Permian-Triassic mass extinction event: 1 2 841 Paleoecology of Lower Triassic carbonates in the western USA. Palaeogeography, 3 4 5 842 Palaeoclimatology, Palaeoecology 116: 1–39. 6 7 8 843 Senowbari-Daryan, B., R. Zühlke, T. Bechstädt, and E. Flügel. 1993. Anisian (Middle Triassic) Buildups 9 10 844 of the Northern Dolomites (Italy): The Recovery of Reef Communities after the Permain/Triassic 11 12 13 845 Crisis. Facies 28: 181–256. 14 15 16 846 Sepkoski, J.J. 1997. Biodiversity: past, present, and future. Journal of Paleontology 71: 533–539. 17 18 19 847 Shigeta, Y., J.D. Zakharov, H. Maeda, A.M. Popov. 2009. The Lower Triassic System in the Abrek Bay 20 21 22 848 Area, South Primorye, Russia. Tokyo: National Museum of Nature and Science. 23 24 25 849 Song, H., P.B. Wignall, Z.Q. Chen, J. Tong, D.P.G. Bond, X. Lai, X. Zhao, H. Jiang, C. Yan, Z. Niu, J. Chen, 26 27 850 H. Yang, Y. Wang. 2011. Recovery tempo and pattern of marine ecosystems after the end-Permian 28 29 30 851 mass extinction. Geology 39: 739–742. 31 32 33 852 Stanley, G.D. 2003. The evolution of modern corals and their early history. Earth-Science Reviews 60: 34 35 853 195-225. 36 37 38 854 Stanley, S.M. 1970. Relation of shell form to life habits of the Bivalvia (Mollusca). Geological Society 39 40 41 855 of America, Memoir 125: 1–296. 42 43 44 856 Stanley, S.M. 2016. Estimates of the magnitudes of major marine mass extinctions in earth history. 45 46 857 Proceedings of the National Academy of Sciences 113: E6325 E6334. 47 – 48 49 50 858 Stiller, F. 2001. Fossilvergesellschaftungen, Paläökologie und paläosynökologische Entwicklung im 51 52 859 Oberen Anis (Mittlere Trias) von Qingyan, insbesondere Bangtoupo, Provinz Guizhou, 53 54 860 Südwestchina. Münstersche Forschung zur Geologie und Paläontologie 92: 1 523. 55 – 56 57 58 59 60 61 62 63 37 64 65 861 Stiller, F., and H. Bucher. 2008. Anisian ammonoids from Qingyan, southwestern China: 1 2 862 biostratigraphical implications for the age of the Qingyan Formation. Swiss Journal of Geosciences 3 4 5 863 101: 547–562. 6 7 8 864 Stur, D.R.J. 1868. Eine Excursion in die Umgegend von St. Cassian. Jahrbuch der Geologischen 9 10 865 Bundesanstalt 18: 529–568. 11 12 13 14 866 Szabó, I., S. Kovács, G. Lelkes, and A. Oravecz-Scheffer. 1979. Stratigraphic investigation of a (triassic) 15 16 867 Pelsonian-Fassanian section at Felsőörs (Balaton Highland, Hungary). Rivista Italiana di 17 18 868 Paleontologia e Stratigrafia 85: 789–806. 19 20 21 22 869 Szulc J. 2007. Sponge-microbial stromatolites and coral-sponge reef recovery in the Triassic of the 23 24 870 western Tethys domain. In The Global Triassic, eds. S.G. Lucas and J.A. Spielmann, 402. New 25 26 871 Mexico Museum of Natural History and Science Bulletin 41. 27 28 29 30 872 Thompson, J.R., E. Petsios, E.H. Davidson, E.M. Erkenbrack, F. Gao, and D.J. Bottjer. 2015. 31 32 873 Reorganisation of sea urchin gene regulatory networks at least 268 million years ago as revealed 33 34 874 by oldest fossil cidaroid echinoid. Scientific Reports 5: 15541. 35 36 37 38 875 Thuy, B., H. Hagdorn, and A.S. Gale. 2017. Paleozoic echinoderm hangovers: Waking up in the 39 40 876 Triassic. Geology 45: 531–534. 41 42 43 877 Tu, C., Z.Q. Chen, and D.A.T. Harper. 2016. Permian-Triassic evolution of the Bivalvia: Extinction- 44 45 46 878 recovery patterns linkes to ecological and taxonomic selectivity. Palaeogeography, 47 48 879 Palaeoclimatology, Palaeoecology 459: 53–62. 49 50 51 880 Twitchett, R.J., and P.B. Wignall. 1996. Trace fossils and the aftermath of the Permo-Triassic mass 52 53 54 881 extinction: evidence from northern Italy. Palaeogeography, Palaeoclimatology, Palaeoecology 55 56 882 124: 137–151. 57 58 59 60 61 62 63 38 64 65 883 Twitchett, R.J., and T. Oji. 2005. Early Triassic recovery of echinoderms. Comptes Rendus Palevol 4: 1 2 884 531–542. 3 4 5 6 885 Urlichs, M. 1977. Zur Altersstellung der Pachycardientuffe und der Unteren Cassianer Schichten in 7 8 886 den Dolomiten (Italien). Mitteilungen der Bayerischen Staatssammlung für Paläontologie und 9 10 887 historische Geologie 17: 15–25. 11 12 13 14 888 Urlichs, M. 1982. Zur Stratigraphie und Fossilführung des Lettenkeupers (Ob. Trias) bei Schwäbisch 15 16 889 Hall (Baden-Württemberg). Jahresberichte und Mitteilungen des Oberrheinischen Geologischen 17 18 890 Vereins, Neue Folge 64: 213–224. 19 20 21 22 891 Urlichs, M. 1992. Vermeintlicher Zwergwuchs bei Muscheln aus dem untersten Trochitenkalk (Oberer 23 24 892 Muschelkalk, Mitteltrias) Württembergs. Carolinea 50: 9–26. 25 26 27 893 Urlichs, M. 1994. Trachyceras LAUBE 1869 (Cephalopoda) aus dem Unterkarn (Obertrias) der 28 29 30 894 Dolomiten (Italien). Stuttgarter Beiträge zur Naturkunde (B) 217: 1–55. 31 32 33 895 Urlichs, M. 2014. A phylogenetic lineage of Myophoria kefersteini subspecies (Bivalvia) from the Late 34 35 896 Ladinian to the Early Carnian (Triassic) in the Dolomites and Julian Alps (Italy). Neues Jahrbuch für 36 37 38 897 Geologie und Paläontologie, Abhandlungen 274: 187–207. 39 40 41 898 Urlichs, M., and Tichy, G. 2000. Zur Alterstellung der Schlernplateau-Schichten (Mitteltrias) am 42 43 899 Schlern (westliche Dolomiten, Italien). Stuttgarter Beiträge zur Naturkunde (B) 297: 1–10. 44 45 46 900 Velledits, F., C. Péró, J. Blau, B. Senowbari-Daryan, S. Kovács, O. Piros, T. Pocsai, H. Szügyi-Simon, P. 47 48 49 901 Dumitricã, and J. Palfy. 2011. The oldest Triassic platform margin reef from the Alpine-Carpathian 50 51 902 region (Aggtelek, NE Hungary): Platform evolution, reefal biota and biostratigraphic framework. 52 53 54 903 Rivista Italiana di Paleontologia e Stratigrafia 117: 221–268. 55 56 57 904 Villier L., A. Brayard, K.G. Bylund, J.F. Jenks, G. Escarguel, N. Olivier, D.A. Stephen, E. Vennin, and E. 58 59 905 Fara. 2018. Superstesaster promissor gen. et sp. nov., a new starfish (Echinodermata, Asteroidea) 60 61 62 63 39 64 65 906 from the Early Triassic of Utah, USA, filling a major gap in the phylogeny of asteroids. Journal of 1 2 907 Systematic Palaeontology 16: 395–415. 3 4 5 6 908 Volz, W. 1896. Die Korallen der Schichten von St. Cassian in Süd-Tirol (Frech, Fritz; Volz, Wilhelm: Die 7 8 909 Korallenfauna der Trias. Monographisch bearbeitet. II). Palaeontographica 43: 1–124. 9 10 11 910 Vörös, A. 2003. The Pelsonian Substage on the Balaton Highland (Middle Triassic, Hungary). 12 13 14 911 Geologica Hungarica 55: 1–52. 15 16 17 912 Waagen, L. 1903. Brachiopoden aus den Pachycardientuffen der Seiser Alpe. Jahrbuch der k. k. 18 19 913 geologischen Reichsanstalt 53: 443–452. 20 21 22 914 Waagen, L. 1907. Die Lamellibranchiaten der Pachycardientuffe der Seiser Alm nebst vergleichend 23 24 25 915 paläontologischen und phylogenetischen Studien. Als Fortsetzung (II. Teil) zu A. Bittner, 26 27 916 Lamellibranchiaten der alpinen Trias. Abhandlungen der k. k. geologischen Reichsanstalt 18: 1– 28 29 30 917 180. 31 32 33 918 Wagner, R. 1897. Beitrag zur genaueren Kenntnis des Muschelkalks bei Jena. Abhandlungen der 34 35 919 Königlich Preussischen Geologischen Landesanstalt Neue Folge 27: 1–105. 36 37 38 920 Weidlich, O., W. Kiessling, and E. Flügel. 2003. Permian-Triassic boundary interval as a model for 39 40 41 921 forcing marine ecosystem collapse by long-term atmospheric oxygen drop. Geology 31: 961–964. 42 43 44 922 Wignall, P.B., and R.J. Twitchett. 2002. Permian-Triassic sedimentology of Jameson Land, East 45 46 923 Greenland: incised submarine channels in an anoxic basin. Journal of the Geological Society 47 48 49 924 London 159: 691–703. 50 51 52 925 Wilckens, R. 1909. Paläontologische Untersuchung triadischer Faunen aus der Umgebung von 53 54 926 Predazzo in Südtirol. Verhandlungen des Naturhistorisch-Medizinischen Vereins Heidelberg 10: 55 56 57 927 81–231. 58 59 60 61 62 63 40 64 65 928 Wöhrmann, S. von, and Koken, E. 1892. Die Fauna der Raibler Schichten vom Schlernplateau. 1 2 929 Zeitschrift der Deutschen geologischen Gesellschaft 44: 167–223. 3 4 5 6 930 Wöhrmann, S. von. 1889. Die Fauna der sogenannten Cardita- und Raibler Schichten in den 7 8 931 Nordtiroler und Bayerischen Alpen. Jahrbuch der k. k. Geologischen Reichsanstalt 39: 181–258. 9 10 11 932 Yang, H., Z.-Q. Chen, Y. Wang, W. Ou, W. Liao, and X. Mei. 2015. Palaeoecology of microconchids 12 13 14 933 from microbialites near the Permian-Triassic boundary in South China. Lethaia 48: 497–508. 15 16 17 934 Yin, H., and E.L. Yochelson. 1983a. Middle Triassic Gastropoda from Qingyan, Guizhou Province, 18 19 935 China: 1. Pleurotomariacea and Murchisoniacea. Journal of Paleontology 57: 162–187. 20 21 22 936 Yin, H., and E.L. Yochelson. 1983b. Middle Triassic Gastropoda from Qingyan, Ghizhou Province, 23 24 25 937 China: 3. Euomphalacea and Loxonematacea. Journal of Paleontology 57: 1098–1127. 26 27 28 938 Yin, H., and E.L. Yochelson. 1983c. Middle Triassic Gastropoda from Qingyan, Guizhou Province, 29 30 939 China: 2. Trochacea and Neritacea. Journal of Paleontology 57: 515 538. 31 – 32 33 34 940 Yose, L.A. 1991. Sequence stratigraphy of mixed carbonate/volcaniclastic slope deposits flanking the 35 36 941 Sciliar (Schlern) – Catinaccio buildup, Dolomites, Italy. In The Northern Margin of the 37 38 942 Schlern/Sciliar-Rosengarten/Catinaccio Platform, eds. R. Brandner, E. Flügel, R. Koch, and L.A. 39 40 41 943 Yose, 17–39. Dolomieu Conference on Carbonate Platforms and Dolomitization, Guidebook 42 43 944 Excursion A, Ortisei (Italy). 44 45 46 945 Zardini, R. 1978. Fossili Cassiani (Trias medio-superiore) Atlante die gasteropodi della formazione di 47 – 48 49 946 S. Cassiano raccolti nella regiona dolomitica attorno a Cortina d’Ampezzo. Cortina d’Ampezzo: 50 51 947 Edizione Ghedina. 52 53 54 948 Zardini, R. 1981. Fossili Cassiani (Trias medio-superiore) Atlante die bivalvi della formazione di S. 55 – 56 57 949 Cassiano raccolti nella regiona dolomitica attorno a Cortina d’Ampezzo. Cortina d’Ampezzo: 58 59 950 Edizione Ghedina. 60 61 62 63 41 64 65 951 Zatoń, M., H. Hagdorn, and T. Borszez. 2013a. Microconchids of the species Microconchus valvatus 1 2 952 (Münster in Goldfuss, 1831) from the Upper Muschelkalk (Middle Triassic) of Germany. 3 4 5 953 Palaeobiodiversity and Palaeoenvironments 94: 453–461. 6 7 8 954 Zatoń, M., P.D. Taylor, and O. Vinn. 2013b. Early Triassic (Spathian) post-extinction Microconchids 9 10 955 from Western Pangea. Journal of Paleontology 87: 159–165. 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 42 64 65 956 Figure captions 1 2 957 3 4 5 6 958 Fig. 1 Locality map showing the position of the studied sections at the Seiser Alm/Alpe di Siusi (red 7 8 959 stars) and the Schlern/Sciliar (blue stars). GPS-coordinates are N 46°30’40.8’’, E 11°33’48.54’’ for 9 10 960 Schlern locality A and N 46°30’36.19’’, E 11°33’51.9’’ for Schlern locality B 11 12 13 14 961 15 16 17 962 Fig. 2 Field photographs. (a) Sampling locality Frommerbach A, showing the investigated section (see 18 19 963 also Fig. 10 [left]). Scale = 1 m. (b) Sampling locality Frommerbach B, showing the investigated 20 21 22 964 section (see also Fig. 10 [right]). (c) Western flank of the Seiser Klamm/Gola di Siusi showing the 23 24 965 Schlernplateau beds (SchPl. beds), the post-volcanic bedded facies of the Rosszähne Fm. (RZ), 25 26 966 volcanic rocks (V) and the pre-volcanic Schlerndolomite of the Rosengarten Fm. (R). The white 27 28 29 967 rectangle indicates the position of (d). (d) Detailed look on the Schlernplateau beds. (e) Bioclastic 30 31 968 rudstone from top of the Schlernplateau beds containing gastropods and bivalves. (f) Oncolites from 32 33 969 the western Schlernplateau. Coin (16 mm) for scale 34 35 36 37 970 38 39 40 971 Fig. 3 Section of the Schlernplateau beds from the western flank of the Seiser Klamm/Gola di Siusi. M 41 42 972 = marl, m = mudstone, w = wackestone, p = packstone, g = grainstone, SP = Schlernplateau. The 43 44 45 973 Ladinian/Carnian boundary lies within bed SP10 46 47 48 974 49 50 51 975 Fig. 4 Rarefaction analyses from the Schlernplateau beds (a) and the Pachycardientuffe (b). Only 52 53 54 976 samples with more than 40 individuals were analysed. See text for details. SP = Schlernplateau, SK = 55 56 977 Seiser Klamm, SK-E = end of Seiser Klamm, FB = Frommmerbach 57 58 59 978 60 61 62 63 43 64 65 979 Fig. 5 Most common bivalves and gastropods collected at the Schlernplateau. (1) Trigonodus costatus 1 2 980 von Wöhrmann and Koken, 1892, left valve interior (1A) and left valve exterior (1B), PZO5984, bed 3 4 5 981 SP7. (2) Cassianella bipartita (von Klipstein, 1845), left valve, PZO5988, bed SP8. (3) Entolioides zitteli 6 7 982 (von Wöhrmann and Koken, 1892), right valve, PZO5986. (4) Myophoriopis richthofeni (Stur, 1868), 8 9 983 left valve, PZO5985, bed SP10. (5) Myophoria kefersteini (Münster in Goldfuss, 1837), left valve, 10 11 12 984 PZO5987, bed SP8. (6) ‘Loxonema’ lineatum von Wöhrmann and Koken, 1892, PZO5994, bed SP8. (7) 13 14 985 ‘Loxonema’ aequale von Wöhrmann and Koken, 1892, PZO5990, bed SP7. (8) Stephanocosmia 15 16 17 986 coronata (von Wöhrmann and Koken, 1892), PZO5989. (9) Zygopleura spinosa von Wöhrmann and 18 19 987 Koken, 1892, PZO5995bed SP8. (10) Pustularia alpina von Eichwald, 1851, PZO5998. (11) Tretospira 20 21 988 multistriata von Wöhrmann and Koken, 1892, PZO5997, bed SP8. (12) Coelostylina solida (von 22 23 24 989 Wöhrmann and Koken, 1892), PZO5996, bed SP7. (13) Neritaria plicatilis (von Klipstein, 1844), 25 26 990 PZO5993, bed SP8. (14) Naticopsis neritacea (Münster, 1841), PZO5991, bed SP7. (15) Palaeonarica 27 28 991 concentrica (Münster, 1841), PZO5992, bed SP7. Scale bars represent 1 cm for bivalves and 0.5 cm 29 30 31 992 for gastropods 32 33 34 993 35 36 37 994 Fig. 6 Bivalves from the Pachycardientuffe. (1) Pachycardia rugosa von Hauer, 1857, right valve 38 39 40 995 exterior (1A) and right valve interior (1B), PZO5980, bed FB B1. (2) Heminajas fissidentata (von 41 42 996 Wöhrmann, 1889), left valve, PZO5983, bed FB A3. (3) Pteria kokeni (von Wöhrmann and Koken, 43 44 997 1892), left valve, PZO5982. (4) Pachycardia plieningeri Broili, 1903, left valve interior (4A) and left 45 46 47 998 valve exterior (4B), PZO5981, bed FB B1. (5) Myoconcha parvula von Wöhrmann and Koken, 1892, 48 49 999 right valve, SMNHS75537. (6) Pteria salomoni (Broili, 1903), right valve, SMNHS75534/1-2. (7) 50 51 1000 Myoconcha curvata Broili, 1903, left valve interior and left valve exterior, SMNHS10274. (8) 52 53 54 1001 Mysidioptera interrupta Broili, 1903, left valve, SMNHS17501/162. (9) Trigonodus rablensis (Gredler, 55 56 1002 1862), right valve exterior (9A) and left valve interior (9B), 10284/1-3. (10) Waagenoperna planata 57 58 59 1003 (Broili, 1903), left valve, SMNHS17501/45. (11) Mysidioptera emiliae Bittner, 1900, right valve, 60 61 1004 SMNHS17501/12. (12) Pteria tofanae (Bittner, 1895), left valve, SMNHS75538, collected at 62 63 44 64 65 1005 Tschapitbach. (13) Newaagia lata (von Klipstein, 1845), right valve interior, SMNHS75536. (14) 1 2 1006 Parallelodon tschapitana (Broili, 1903), right valve, SMNHS17501/250. Scale bars represent 1 cm 3 4 5 6 1007 7 8 9 1008 Fig. 7 Q-mode (samples) and R-mode (species) cluster analysis of samples with more than 40 10 11 1009 individuals from the Schlernplateau beds using unweighted pair-group average and Morisita 12 13 14 1010 similarity index. The circle sizes represent the abundance of species within the corresponding sample 15 16 1011 (small circle = 1 individual, small medium-sized circle = 2-5 individuals, big medium-sized circle = 6-20 17 18 1012 individuals, big circle = more than 20 individuals). SP = Schlernplateau, SK = Seiser Klamm, SK-E = end 19 20 21 1013 of Seiser Klamm. See text for details 22 23 24 1014 25 26 27 1015 Fig. 8 Rank-abundance distributions and guild-structures of the entire fauna of the Schlernplateau 28 29 30 1016 beds (a) and Pachycardientuffe (b). n = total number of individuals 31 32 33 1017 34 35 36 1018 Fig. 9 Rank-abundance distributions and guild-structures of representative fossiliferous beds SP14 37 38 1019 (a), SP7 (b) and SP3 (c) from the Schlernplateau beds (see also Fig. 3). n = total number of individuals 39 40 41 42 1020 43 44 45 1021 Fig. 10 Sections Frommerbach A and B of the Pachycardientuffe from the Seiser Alm/Alpe di Siusi 46 47 48 1022 49 50 51 1023 Fig. 11 Rank-abundance distributions and guild-structures from the fossiliferous beds FB A3 (a), FB B2 52 53 54 1024 (b) and FB B1 (c) from the Pachycardientuffe (see also Fig. 10). n = total number of individuals 55 56 57 1025 58 59 60 61 62 63 45 64 65 1026 Fig. 12 Lithological samples from locality Frommerbach A showing the highly fossiliferous limestone 1 2 1027 (a) and volcanoclastic sandy siltstone (b-c) containing disarticulated, randomly arranged bivalves. 3 4 5 1028 Scales are 23 mm (diameter of the coin) for (a) and 5 cm for (b) and (c) 6 7 8 1029 9 10 11 1030 Fig. 13 (a) Palaeoenvironmental reconstruction of the Schlern and Seiser Alm area during the Late 12 13 14 1031 Ladinian showing the distribution of sedimentary facies and the potential source areas of the 15 16 1032 Pachycardientuffe. (b) Panorama image taken from the top of the eastern margin of the Schlern 17 18 1033 mountain, showing present-day position of the former lagoon (Schlernplateau beds, 1), the post- 19 20 21 1034 volcanic reef (Rosszähne Fm., 2), the probable position of the suggested volcanic island (3) and the 22 23 1035 basin (Seiser Alm basin, Pachycardientuffe and Wengen Fm., 4) 24 25 26 1036 27 28 29 30 1037 Fig. 14 Diversity changes throughout the Early and Middle Triassic based on data shown in Tab. 1; see 31 32 1038 text for further explanation. (a) Total number of species. (b) Gastropod/bivalve-ratio. Blue lines mark 33 34 1039 formations with carbonate platform facies. G = Griesbachian; D = Dienerian; Sm = Smithian; Sp = 35 36 37 1040 Spathian, A = Aegean; B = Bithynian; P = Pelsonian; I = Illyrian; F = Fassanian; L = Longobardian; J = 38 39 1041 Julian; Carn. = Carnian; LT = Late Triassic. Marmolada = Marmolada Limestone; PachyT = 40 41 1042 Pachycardientuffe; Latemar = Latemar Limestone; SchPl = Schlernplateau beds 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 46 64 65 1043 Table captions 1 2 1044 3 4 5 6 1045 Tab. 1 Benthic species richness of selected Early and Middle Triassic faunas. B = bivalves, G = 7 8 1046 gastropods, Br = brachiopods, C = corals, Sp = sponges, Cr = crinoids, E = echinoids, A = asteroids, Bry 9 10 1047 = bryozoans, Cru = crustaceans, H = holothurioids, M = microconchids, O = ophioroids, Pc = 11 12 13 1048 polychaete worms, Pp = polyplacophorans, Sc = scaphopods, ? = these taxa were reported but the 14 15 1049 exact number of species is unknown. Information about the age of the lithological units come from 16 17 1050 Fürsich and Wendt 1977, Broglio Loriga et al. 1990, Brack and Rieber 1993, Brack et al. 1996, Vörös 18 19 20 1051 2003, Manfrin et al. 2005, Stiller and Bucher 2008, Hagdorn and Nitsch 2009, Shigeta et al. 2009, 21 22 1052 Balini et al. 2010, Hofmann et al. 2013a, 2013b, 2014, 2015, Urlichs 2014, Foster et al. 2015, 2017, 23 24 25 1053 Hausmann and Nützel 2015, Hautmann et al. 2015, Brayard et al. 2017, Foster and Sebe 2017. See 26 27 1054 text for details 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 47 64 65 1055 Online Resources captions 1 2 1056 3 4 5 6 1057 Online Resource 1 Excel tables showing the quantitative matrix of fossil occurrences used for data 7 8 1058 analyses, the comparison between the faunas of the Schlernplateau beds and the Pachycardientuffe 9 10 1059 as well as the number of species, number of individuals and Simpson-D values for each fossiliferous 11 12 13 1060 bed 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 48 64 65 Revision notes Click here to download Authors' Response to Reviewers' Comments Revision notes 2.docx

Revision notes

Ref.: Ms. No. PAZE-D-17-00052R1

Palaeoecology of Late Ladinian (Middle Triassic) benthic faunas from the Schlern/Sciliar and Seiser Alm/Alpe di Siusi area (South Tyrol, Italy)

General note: Please see attached document with “track changes” for the details of our changes in response to the reviewer’s comments.

Reviewer’s comments

AUTHOR AFFILIATIONS:

Once again, to avoid any confusion, please please split the affiliations of Alexander Nützel ... SNSB- BSPG and LMU are clearly different institutions (see my former comments)

Response: We split the affiliation of Alexander Nützel and apologize that we did not do that in the first place.

TEXT:

Page 22, line 505 ... Szulc INSTEAD OF Sculz 2007

Page 23, line 524 ... Kozur and Mostler 1973 IS MISSING in your REFERENCES

Page 23, lines 524/525 ... Mostler 1977 IS MISSING in your REFERENCES

Page 23, line 526 ... Reich 2013 IS MISSING in your REFERENCES

Page 24, line 548 ... Gaetani et al. 1981 INSTEAD OF Gaetani 1981

Page 28, line 646 ... Eichwald, E. von. 1851. INSTEAD OF Eichwald, M. 1851.

Page 28, line 646 ... Geognosie, auf INSTEAD OF Geognosie auf Page 28, line 647 ... die Eifel, Tyrol, Italien, Sizilien INSTEAD OF die Eife, Tirol, Sizilien

Page 30, line 691 ... Gümbel, C.W. 1861. INSTEAD OF Gümbel, C.W. von 1861. ...(Gümbel was nobled in 1882)

Page 33, lines 754/55 ... implement Kozur and Mostler 1973 (see Page 23, line 524)

Page 34, lines 782/783 ... implement Mostler 1977 (see Page 23, lines 524/525)

Page 36, lines 820/821 ... implement Reich 2013 (see Page 23, line 526)

Page 36, lines 825-826 ... DELETE Sano and Nakashima 1997 which is not cited in text etc.

Page 39, line 887 ... "Trachyceras" IN ITALICS please

Page 39, line 889 ... "Myophoria kefersteini" IN ITALICS please

Page 40, line 928 ... Yin INSTEAD OF Ying

Page 41, line 930 ... Yin INSTEAD OF Ying

Page 41, line 932 ... Yin INSTEAD OF Ying

Page 43 ... line 981 ... von Eichwald, 1851 INSTEAD OF Eichwald, 1851

Response: As requested we changed the above mentioned text passages and deleted the work of Sano and Nakazawa (1997). Naturally, we added the work of Kozur and Mostler (1973), Mostler (1977) and Reich (2013) to the reference list and apologize that this mistake did not track our attention before.

REFERENCES:

Please check again my former comments (comment 5) ... Please use an en-dash NOT a hyphen netween page numbers

Response: All hyphens between page numbers were changed into en-dashes.

TABLE:

Frizzell and Exline (1956) INSTEAD OF Frizzell and Exline (1955)

Response: We changed the above mentioned text passage as requested.

ONLINE RESOURCE:

Please re-format your species names ... "sp.", "sp. 1...", "cf.", "n.sp." NOT IN ITALICS

Response: We performed the necessary changes in our electronic supplementary material.

Further changes

Independently from the reviewer’s suggestions we performed some minor changes on the manuscript, which can be seen in the track-edited version of the manuscript. Track-edited version Click here to download Authors' Response to Reviewers' Comments Manuscript track-edited version 2.docx

1 Palaeoecology of Late Ladinian (Middle Triassic) benthic faunas from the Schlern/Sciliar and Seiser

2 Alm/Alpe di Siusi area (South Tyrol, Italy)

3

4 Evelyn Friesenbichler1*, Michael Hautmann1, Alexander Nützel2, Max Urlichs3, Hugo Bucher1

5

6 1 Paläontologisches Institut und Museum, University of Zurich, Karl-Schmid-Strasse 4, 8006 Zurich,

7 Switzerland

8 2 SNSB - Bayerische Staatssammlung für Paläontologie und Geologie, Richard-Wagner-Straße 10,

9 80333 Munich, Germany and Department of Earth and Environmental Sciences, Palaeontology &

10 Geobiology, Geo-Bio-Center LMU, Richard-Wagner-Straße 10, 80333 Munich, Germany

11 3 Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany

12

13 * corresponding author; e-mail: [email protected]; tel: +41 44 634 23 47

14

15 Acknowledgements

16

17 This study was supported by the Swiss National Science Foundation project 200021-162402/1 (to

18 M. Hautmann). Markus Hebeisen is thanked for his support during the preparation of fossil material

19 and Rosi Roth is thanked for helping during photographing the fossils. Michael Rasser (State Museum

20 of Natural History Stuttgart, Germany) kindly provided access to the collection of the Stuttgart State

21 Museum of Natural History and lending some specimens of the Stuttgart collection. This article

1

22 benefited from careful reviews by William Foster, Christopher A. McRoberts, Simon Schneider and

23 Mike Reich.

2

24 Abstract

25

26 The Schlern and Seiser Alm area (South Tyrol, Italy) is a classical locality for studies of Middle

27 Triassic platform to basin transitions, yet details of the palaeoecology of the rich benthic faunas of

28 this area have been insufficiently known. We herein present a detailed palaeoecological study of the

29 fauna from the Schlernplateau beds (late Ladinian to early Carnian) and the more or less time-

30 equivalent Pachycardientuffe (late Ladinian), which is based on quantitative faunal data. The

31 palaeoecology as well as sedimentary features suggest that the fauna of the Schlernplateau beds

32 represents a lagoonal soft bottom fauna. The high species richness of the fauna as well as the locally

33 restricted occurrences of fossils indicate an open lagoonal setting palaeogeographically close to an

34 oceanic inlet. The high evenness of the fauna is probably a result of time-averaging. In contrast, the

35 fauna of the Pachycardientuffe shows clear indications of transportation. Ecological features of this

36 fauna and palaeogeographic reconstructions suggest three potential source areas: (1) the lagoon

37 represented by the Schlernplateau beds, (2) the reef fringing this lagoon and (3) a shallow clastic

38 coast of a nearby volcanic island and/or submarine high. A comparison between diversities of

39 selected Early and Middle Triassic lithological units revealed the increasing species richness of all

40 major benthic taxa during the Middle Triassic and a shift from bivalve dominated Early Triassic faunas

41 to gastropod dominated faunas.

42

43 Keywords: Middle Triassic, biotic recovery, quantitative faunal analyses, palaeoecology,

44 palaeoenvironment.

45

46 Kurzfassung

47

3

48 Obwohl das Gebiet des Schlern und der Seiser Alm (Südtirol, Italien) seit langem als klassische

49 Lokalität für die Erforschung mitteltriassischer Plattform-Becken-Übergänge gilt, ist die Paläoökologie

50 ihrer reichen benthischen Fauna unzureichend erforscht. Wir präsentieren eine detaillierte

51 paläoökologische Studie der Fauna der Schlernplateau-Schichten (spätes Ladinium bis frühes

52 Karnium) und der mehr oder weniger gleichalten Pachycardientuffe (spätes Ladinium), welche auf

53 quantitativen Daten basiert. Die Paläoökologie sowie der sedimentologische Kontext legen nahe,

54 dass die Fauna der Schlernplateau-Schichten eine lagunäre Weichbodenfauna repräsentiert. Ihre

55 hohe Diversität sowie die lokal begrenzten Vorkommen der Fossilien deuten auf eine offene Lagune

56 hin, die sich paläogeographisch nahe an einem Einlass zum offenen Meer befand. Die hohe

57 Gleichmäßigkeit der Fauna ist höchstwahrscheinlich das Resultat von „time-averaging“. Im Gegensatz

58 dazu gibt es in der Fauna der Pachycardientuffe klare Hinweise auf Transport. Die Ökologie dieser

59 Fauna und paläogeographische Rekonstruktionen lassen auf drei potentielle Herkunftsgebiete

60 schließen: (1) die Lagune, welche durch die Schlernplateau-Schichten repräsentiert wird, (2) das

61 Saumriff dieser Lagune und (3) eine flache, siliziklastische Küste nahe einer vulkanischen Insel

62 und/oder einer submarinen Hochzone. Ein Vergleich der Diversitäten ausgewählter früh- und

63 mitteltriassischer lithologischer Einheiten offenbart einen Anstieg der Diversität während der Früh-

64 und Mitteltrias sowie einen Wechsel von Muschel-dominierten frühtriassischen Faunen zu

65 Gastropoden-dominierten Faunen während der Mitteltrias.

66

67 Schlüsselwörter: Mitteltrias, ökologische Erholung, quantitative Faunen-Analysen, Paläoökologie,

68 Paläo-Umwelt.

69

70 1. Introduction

71

4

72 The recovery from the end-Permian mass extinction, which killed between 81% (Stanley 2016)

73 and 96% (Raup 1979) of marine species, provides a unique test-case for studying long-term

74 evolutionary responses to a sudden and massive loss of species on a global scale. Scientific interest in

75 studying the recovery pattern is twofold: it allows testing macroevolutionary models (e.g., Brayard et

76 al. 2009; Hautmann et al. 2015), and it represents a (pessimistic) analogue for the long-term future of

77 biosphere in the face of anthropogenically caused environmental changes (Payne and Clapham

78 2012).

79 Understanding evolutionary processes requires the study of species in their ecological context.

80 Concerning the recovery from the end-Permian mass extinction, much palaeoecological work has

81 been done in the past decades, but these studies almost exclusively concentrated on the Early

82 Triassic (e.g., Schubert and Bottjer 1995; Twitchett and Wignall 1996; Hofmann et al. 2013a, 2013b,

83 2014, 2015; Hautmann et al. 2011, 2015; Foster et al. 2017), which is the ca. 4 Ma time interval that

84 followed the end-Permian mass extinction. However, biodiversity in terms of global taxonomic

85 richness did not reach the pre-extinction level before the Late Jurassic (Sepkoski 1997) or later (Alroy

86 et al. 2008). With regard to alpha-diversity in local environments, the major recovery step seems to

87 have taken place during the Middle Triassic, as evidenced by the striking differences between most

88 diverse Early Triassic communities that include around 45 benthic species (Hautmann et al. 2013,

89 2015; Hofmann et al. 2013b, 2014) and, e.g., the hyperdiversified fauna from the Longobardian–

90 Julian (Late Ladinian–Early Carnian) Cassian Fm. of northern Italy (>1000 benthic species; Tab. 1). The

91 progress in recovery during the Middle Triassic has been addressed in several recent studies (e.g.,

92 Payne 2005; Payne et al. 2006; Song et al. 2011; Velledits et al. 2011; Foster and Sebe 2017) but data

93 are still relatively scarce.

94 This study is based on quantitative bed-rock sampling in the area of Schlern/Sciliar mountain and

95 Seiser Alm/Alpe di Siusi (South Tyrol, Italy; Fig. 1), which is a paradigm for a Middle to Late Triassic

96 reef-to-basin landscape. It is one of the few places in the world where the platform-slope-basin

97 transitional zone is preserved in its primary setting (Keim 2008; Brandner et al. 2016), giving the 5

98 opportunity to study the ecology of concurrent faunas in different environments. Sampling was

99 carried out (1) in sediments of a lagoonal setting represented by the locally highly fossiliferous

100 Schlernplateau beds on the Schlern mountain and (2) in the basinal setting of the ‘Pachycardientuffe’

101 at the Seiser Alm, which represent approximately the same stratigraphic age (see chapter 3). The

102 palaeoecology and species distributions of both faunas are discussed in the context of their

103 depositional environments, and their diversities are compared with published data on Early–Middle

104 Triassic benthic communities from other regions, which allows preliminary insights into diversity

105 changes through the Early and Middle Triassic time interval.

106

107 2. Material and Methods

108

109 Fieldwork was carried out in August 2016 at the Seiser Alm/Alpe di Siusi and the Schlern/Sciliar

110 mountain in South Tyrol/Alto Adige in northern Italy. At the Seiser Alm, fossils were sampled in the

111 gorge of the Frommerbach/Rio Frommer and one of its eastern tributaries (Figs 1 and 2a-b). On the

112 Schlern, fossils were mainly collected on the western flank of the Seiser Klamm/Gola di Siusi (Figs 1

113 and 2c-d). Outcropping rock material of fossiliferous beds was quantitatively sampled until the

114 occurrences of fossils were exhausted. Additionally, fossils quantitatively collected by M. Urlichs

115 between 1998 and 2012 at the same localities were included in the analyses. Specimens used for the

116 palaeoecological analyses are housed at the State Museum of Natural History Stuttgart (Germany,

117 inventory numbers SMNHS17501/250 - SMNHS75538) and the Museum of Nature South Tyrol (Italy,

118 inventory numbers PZO5980 - PZO5998). For taxonomic identification fossils were first washed and

119 then, when necessary, prepared with micro jacks. A quantitative matrix of fossil occurrences was

120 compiled at species level (Online Resource 1). For bivalves, the number of individuals has been

121 inferred by counting only the most numerous valves (left or right) together with articulated

122 specimens. Only fossils that (1) were clearly relatable to specific beds of the investigated outcrops 6

123 and (2) samples with more than 40 individuals were used for palaeoecological data analyses.

124 Sedimentological observations in the field and information from ecological analyses were used for

125 the interpretation of the depositional environment. Our complete dataset comprises 2723 individuals

126 assigned to 53 species for the Schlernplateau beds and 651 individuals assigned to 59 species for the

127 Pachycardientuffe.

128 Ecological parameters and statistical analyses were calculated using the software PAST, version

129 2.17 (Hammer et al. 2001). Based on the species-abundance data, rarefaction analyses were carried

130 out for each sample in order to compare species richness of faunas with different sample sizes and to

131 assess the relative completeness of sampling. For further ecological characterization, rank-

132 abundance distributions, trophic nuclei, and ecological pie charts were used. The trophic nucleus of

133 an association comprises those species that contribute 80% of the total number of individuals in an

134 association (Neyman 1967). Following Aberhan (1994), species were assigned to 10 trophic guilds,

135 based on functional morphology, mode of life of related Recent species, information from the

136 literature, and the sedimentological context. Gastropods were collectively assigned to epifaunal

137 herbivores and/or detritus-feeders, since evidence for more specialized feeding lacks. For the

138 identification of recurrent assemblages and groups of ecologically related species, Q-mode cluster

139 analyses were calculated for samples containing more than 40 individuals, using the Morisita

140 similarity index (Morisita 1959) and unweighted pair-group average. Additionally, R-mode cluster

141 analyses were used for the identification of sets of species that typically co-occur.

142

143 3. Geologic setting

144

145 The Schlern and the adjacent Rosengarten/Catinaccio represent a Middle Triassic carbonate

146 platform with northeastward, eastward to southeastward inclined reef slopes (Brandner 1991;

7

147 Bosellini 1998; Brandner et al. 2007; Keim 2008). Towards the west, there was probably a connection

148 to the Mendel platform (Keim 2008). The Schlern Mountain is build up by (1) the pre-volcanic, up to

149 800 m thick ‘Schlerndolomite’ (Rosengarten Fm.), (2) up to 40 m thick volcanic rocks and (3) post-

150 volcanic, well-bedded dolomite formerly known as ‘Rosetta Dolomite’ or ‘Cassian Dolomite’, now

151 interpreted as the platform facies of the Rosszähne Fm. The latter is overlain by (4) the

152 Schlernplateau beds, which is a sequence of carbonate rocks and grey to reddish marls that are

153 locally highly fossiliferous and commonly interpreted as lagoonal sediments (Figs 2c-d; Keim 2008;

154 Brandner et al. 2016). Findings of the biostratigraphically important ammonoid Prototrachyceras

155 archelaus (Laube, 1869b) indicate a Late Ladinian age of the Schlernplateau beds (Brandner 1991;

156 Urlichs and Tichy 2000). However, the occurrence of the bivalve Myophoria kefersteini kefersteini

157 (Münster in Goldfuss, 1837) in the uppermost 6 m of the Schlernplateau beds suggests that the top

158 of the formation extends into the Lower Carnian (Urlichs 2014). The thickness of the Schlernplateau

159 beds varies from only a few meters to a maximum of ca. 40 m. Towards the platform margins, the

160 Schlernplateau beds interfinger with the reef flat facies.

161 During deposition of the Rosszähne Fm. and Schlernplateau beds at the top of the Schlern

162 carbonate platform, the adjacent Seiser Alm basin was filled with a more than 500 m thick

163 successions of volcanoclastic detritus of the Wengen Fm. and, later, the mixed siliciclastic-carbonate

164 succession of the St. Cassian/San Cassiano Fm. (Brandner 1991). Within the Wengen Fm. of the Seiser

165 Alm, the Late Ladinian (Urlichs 1977, 1994, 2014) ‘Pachycardientuffe’, which are named for the mass

166 occurrence of the bivalve Pachycardia rugosa von Hauer, 1857, represent several (at least two)

167 conglomeratic, highly fossiliferous intercalations that have been mapped as distal parts of the

168 ‘Marmolada Conglomerate’ in the geological map of the western Dolomites (Autonome Provinz

169 Bozen (ed.) 2007). However, the Marmolada Conglomerate in the strict sense is a volcanogenic

170 conglomerate that was transported from a source area near the Marmolada Mountain (ca. 20 km

171 east of the study area) into adjacent basins (e.g., Bosellini 1998; Bottoli and Trombetta 1998). In

172 contrast, the lithology of the Pachycardientuffe is dominated by non-dolomitized limestone pebbles

8

173 and cobbles (small to medium sized ‘Cipit boulders’) that originate from adjacent carbonate

174 platforms. We therefore assume that the Pachycardientuffe represent distal equivalents of tongues

175 of foreslope talus deposits, as described by Yose (1991) in a proximal facies at the Mahlknecht cliff,

176 around 4 km SE of our sampling localities. The admixed volcanogenic pebbles and volcanoclastic

177 matrix could either have been taken up during transportation, or stem from a second source area, or

178 could represent the result of mixing of carbonate and volcanogenic components by reworking (see

179 chapter 5.2.).

180

181 4. The Schlernplateau beds

182

183 Early workers erroneously correlated the fauna of the Schlernplateau beds with the younger Raibl

184 Fm. (Carnian), based among other reasons on the occurrence of Myophoria kefersteini (Münster in

185 Goldfuss, 1837; e.g., von Richthofen 1860). This view was also shared by von Wöhrmann and Koken

186 (1892) in their comprehensive description of the Schlernplateau fauna and was maintained by most

187 subsequent authors until Brandner (1991), Urlichs and Tichy (2000) and Urlichs (2014) provided

188 evidence for a Late Ladinian (Prototrachyceras archelarus–Frankites regoledanus ammonoid zone) to

189 early Carnian (Daxatina canadensis ammonoid zone) age. In addition to ammonoid finds, the key

190 observation for the revised stratigraphy was the existence of several subspecies of Myophoria

191 kefersteini, which are indicative of different stratigraphic intervals (Urlichs 2014). With this updated

192 stratigraphic information, Schlernplateau beds and Pachycardientuffe are more or less time-

193 equivalent units.

194 Von Wöhrmann and Koken (1892) described 72 species, including 33 gastropod species, 24

195 bivalve species, eight cephalopod species, three echinoid species, two coral species and one crinoid

196 species from the Schlernplateau beds. Notably, nearly all specimens of this rich fauna stem from a

9

197 limited area at the western flank and southern part of the Seiser Klamm. The only other locality that

198 is explicitly mentioned in von Wöhrmann and Koken (1892) is the ‘Burgstall’, some hundred meters

199 east of the Seiser Klamm, where one coral species (Thecosmilia rothpletzi von Wöhrmann and Koken,

200 1892), fragments of crinoids and echinoid spines were found. Our survey confirms that fossils are

201 scarce or absent at other places of the Schlernplateau. At the Seiser Klamm locality, fossils are most

202 abundant in the upper part of the section, particularly in several layers of red marl (Fig. 3). Fossils are

203 also abundant in several rudstone beds near the top of the section (Fig. 2e), but the crystallized

204 limestone makes it difficult to extract them from the matrix.

205 A detailed section was measured at the western flank of the Seiser Klamm and samples were

206 taken from three localities on the Maultierrücken/Dorsale del Mul (Figs 1 and 3). The investigated

207 section starts with an alternation (ca. 20 m) of partly dolomitic limestone beds, which are separated

208 by marls or covered intervals that probably also represent fine-grained sediments. The limestone and

209 dolomite beds are mud-, wacke-, pack- and grainstones that sometimes contain bioclasts like bivalve

210 and gastropod shells and, less common, ooids. The overlaying marl beds (SP3 and SP5) are highly

211 fossiliferous. Above these marls, an around 3 m thick sequence of thick-bedded wackestone forms a

212 prominent cliff, the top of which is formed by a 30 cm thick oolitic limestone bed (at a height of 25

213 m). Several marly, highly fossiliferous intervals above this cliff provided the majority of fossils studied

214 herein. Within the marly intervals, laterally discontinuous oolithic limestone beds are repeatedly

215 intercalated, which sometimes contain bioclasts such as gastropod and bivalve shells. A notable

216 lithological marker bed near the top of the section contains shell debris, ooids, and interspersed

217 pisolithic iron ore grains. Surface ornamentations on the fossils are generally well preserved. This lack

218 of abrasion as well as the fact that some bivalves were preserved with conjoined valves indicate that

219 the fossils were not transported over larger distances. According to Urlichs and Tichy (2000) and

220 Urlichs (2014) the uppermost layers of the investigated section are already Carnian in age.

221 Accordingly, the Ladinian/Carnian boundary lies within fossiliferous bed SP10 (Fig. 3).

222 10

223 4.1. Fauna of the Schlernplateau beds

224

225 A total number of 53 species occurring in 16 fossiliferous beds (SP1-SP16, Fig. 3) was identified,

226 including 27 gastropod species, 19 bivalve species, four coral species and one sponge species (Online

227 Resource 1). Notably, brachiopods are absent, and only one crinoid columnal fragment and one

228 echinoid spine were found. In terms of species richness, our collection is comparable to the diversity

229 described in the monograph of von Wöhrmann and Koken (1892; 72 species). Species richness within

230 the different beds varies between 2 to 32. However, rarefaction analyses of samples from each fossil

231 horizon indicate different degrees of sampling completeness (Fig. 4a).

232 The 10 most frequent species are, in descending order, Myophoria kefersteini (Münster in

233 Goldfuss, 1837) (bivalve, 16.3%), Heminajas fissidentata (von Wöhrmann, 1889) (bivalve, 11.4%),

234 Pachycardia rugosa von Hauer, 1857 (bivalve, 9.7%), Neritaria plicatilis (von Klipstein, 1844)

235 (gastropod, 8.3%), Naticopsis neritacea (Münster, 1841) (gastropod, 8%), Coelostylina solida (von

236 Wöhrmann and Koken, 1892) (gastropod, 5.8%), Palaeonarica concentrica (Münster, 1841)

237 (gastropod, 4.3%), Myophoriopis richthofeni (Stur, 1868) (bivalve, 4%), Trigonodus rablensis (Gredler,

238 1862) (bivalve, 3.8%) and ‘Loxonema’ aequale von Wöhrmann and Koken, 1892 (gastropod, 3.3%).

239 These 10 species together with Cassianella bipartita (von Klipstein, 1845) (bivalve, 3.2%) and

240 Stephanocosmia coronata (von Wöhrmann and Koken, 1892) (gastropod, 3%; Fig. 5 and 6) constitute

241 the trophic nucleus of the complete fauna at the Schlernplateau. Simpson’s index of dominance of

242 associations from different fossiliferous beds is generally high and varies between 0.06 and 0.30, with

243 an average value of 0.2 (Online Resource 1).

244 Cluster analysis (Fig. 7) did not reveal the existence of clearly distinct associations. However, a

245 cluster of samples that are dominated by Myophoria kefersteini is set off from samples in which

246 species of Pachycardia and Trigonodus are more abundant. Because Myophoria, Trigonodus and

247 Pachycardia are not only phylogenetically closely related but also ecologically similar (both are

11

248 shallow infaunal suspension feeders), we do not believe that these two clusters represent

249 fundamentally different environmental conditions. As an alternative explanation, it seems possible

250 that haphazardness of larvae settlement had randomly favoured different dominant species at

251 different times.

252 Although the fauna of the Schlernplateau beds is gastropod dominated in terms of species

253 richness, it is bivalve dominated in terms of total number of individuals. Accordingly, most species

254 were epifaunal herbivores and/or detritus-feeders, whereas almost half of all collected individuals

255 are shallow infaunal suspension-feeders, followed by epifaunal herbivores and/or detritus-feeders

256 and free-lying epifaunal suspension-feeders (Fig. 8a). However, the guild-specific diversity and

257 absolute abundance of trophic guilds can vary between different fossiliferous beds (Fig. 9).

258 Frequency distributions of assemblages from different fossiliferous beds show different patterns,

259 but a remarkable aspect is that many show an unusually high degree of evenness (Figs 9b-c). The lack

260 of abrasion, occurrence of bivalves with conjoined valves, and the sedimentological context

261 (occurrence in fine-grained sediments) exclude that this pattern is due to effects of faunal mixing by

262 transportation. Another notable aspect is that there are often lateral differences in the faunal

263 composition within single fossiliferous beds (Fig. 9).

264

265 4.2. Interpretation of the Schlernplateau fauna

266

267 The geologic context leaves little doubt that the Schlernplateau beds represent lagoonal

268 sediments of the platform interior. This is evident by the sedimentary features of the Schlernplateau

269 beds themselves, which include the coexistence of high- and low-energy sediments (e.g., oolites and

270 shell coquinas (rudstones) versus marls, wackestones, and early diagenetic dolomites) and the

271 presence of locally abundant oncoids (Fig. 2f). It is further corroborated by the lateral facies change,

272 which includes the transition to fringing coral reefs on the supposed seaward side and to pedogenic 12

273 sediments at places where the platform was emerged (Brandner et al. 2016 and own observations).

274 Finally, the ecology of the documented fauna is in accordance with a lagoonal setting because

275 shallow infaunal suspension-feeders and free-lying epifaunal suspension-feeders dominate, which

276 are commonly associated with a soft substratum.

277 Given the lagoonal setting, two aspects of the Schlernplateau fauna are remarkable. First, the

278 species richness is unusually high for this environment. Although decidedly stenohaline organisms

279 lack (e.g., brachiopods) or are extremely rare (e.g., echinoderms and ammonoids), the high species

280 richness excludes that the lagoon was (at the place where the samples were taken) affected by high

281 environmental stress such as changing salinity, which often characterizes restricted lagoons (e.g.,

282 Fürsich 1994; Fürsich et al. 1995). The second unusual aspect is the extremely high evenness in many

283 samples. Based on the state of preservation and the occurrence in marly beds (see above), faunal

284 mixing by transportation can be excluded. However, no examples of primary causes for such high

285 evenness are known to us from ecologically comparable faunas. We therefore assume that the high

286 evenness may, at least partly, reflect the effects of time-averaging. Fürsich and Aberhan (1990) nicely

287 demonstrated how time-averaging may produce abundance patterns similar to those that we

288 observed in the Schlernplateau beds.

289 Our summary interpretation of the sedimentological and faunistic data is that the Schlernplateau

290 beds represent an open lagoon and that the outcrop area around the Seiser Klamm was

291 palaeogeographically close to an oceanic inlet that prevented large-scale fluctuations in salinity and

292 temperature, provided supply of food for filter feeding organisms, and was the source of larvae that

293 settled within the lagoon. The randomness in the composition of larvae that reached the lagoon may

294 be partly responsible for differences in the faunal composition. In this model, successive

295 communities went back to different settlement events, each with a particular taxonomic composition

296 of larvae and thus resulting in different dominance of the resulting cohort. Superimposition of

297 abundance patterns of successive communities by time-averaging then caused a pattern where many

298 species are nearly equally abundant. However, prevalent species probably dominated at different 13

299 times. We note that time-averaging might have also contributed to the high overall species richness,

300 but even if this effect is considered, the Schlernplateau beds are still remarkably species rich.

301 Additional support for the open lagoon model comes from the fact that fossil occurrences are

302 locally restricted (close to oceanic inlets in our interpretation). From a sedimentological perspective,

303 the laterally discontinuous oolitic limestone beds that are intercalated in the low-energy marls

304 probably represent washover deposits from a nearby barrier, which however was discontinuous near

305 the fossil locality.

306

307 5. The Pachycardientuffe/Tufi a Pachycardie

308

309 Like the fauna from the Schlernplateau beds, the highly diverse fauna of the Pachycardientuffe is

310 known for more than 100 years. Broili (1903) described 157 species, including 122 bivalve species, 18

311 brachiopod species, 13 echinoid species and four crinoid species. Blaschke (1905) added the

312 description of 77 gastropod species, as well as three sponge species, five coral species, one

313 scaphopod species and 14 bivalve species that were not described in Broili (1903). A detailed

314 description of brachiopods from the Pachycardientuffe was given by Waagen (1903). After a revision

315 by Waagen (1907) a total number of 165 bivalve species is described from the Pachycardientuffe.

316 Overall, more than 270 species have been reported from the Pachycardientuffe thus far (Tab. 1). It

317 should be noted, however, that some of the major fossil localities (e.g., at the Tschapitbach/Rio Cipit)

318 are currently covered and therefore not accessible.

319 The investigated section at locality Frommerbach A is located in the gorge of the Frommerbach

320 itself (Figs 1, 2a and 10). It starts with a fossiliferous conglomerate at 1850 m altitude that overlies

321 volcanoclastic sandstone beds of the Wengen Fm. with an erosional disconformity. The pebbles are

322 mainly limestones and less commonly rounded volcanic rocks, which are embedded in a siltstone

14

323 matrix. The overlaying bedded sandstone contains limestone blocks at its top. The components of

324 the following fossiliferous conglomerate consist of limestone and volcanoclastics. After a 3 m thick

325 interval of siltstone, highly fossiliferous sandy siltstones follow on top of this section. The

326 investigated section Frommerbach B is located in an eastern tributary of the Frommerbach (Figs 1, 2b

327 and 10) at 1850 m altitude. The rocks are matrix supported, fossiliferous, pebble bearing,

328 volcanogenic, middle- to coarse grained sandstones. As pointed out by Urlichs (2014), the

329 fossiliferous rocks in the two Frommerbach sections do not belong to the same horizon. The

330 investigated rocks of section Frommerbach A are younger than the rocks from Frommerbach B. For a

331 detailed stratigraphic correlation between the sections see Fig. 3 in Urlichs (2014).

332

333 5.1. Fauna of the Pachycardientuffe

334

335 At the two Frommerbach localities a total number of 59 species was found, which is much less

336 than the overall species richness of the Pachycardientuffe (277 species), as described in Broili (1903),

337 Waagen (1903), Blaschke (1905) and Waagen (1907). These 59 species include 30 bivalve species, 16

338 gastropod species, five echinoid species, three coral species, three crinoid species and two

339 brachiopod species. Sponges are absent. The seven most frequent species are, in descending order,

340 Pachycardia rugosa von Hauer, 1857 (54.4%), Pachycardia plieningeri Broili, 1903 (27.3%),

341 Trigonodus rablensis (Gredler, 1862) (3.7%), Myophoria kefersteini (Münster in Goldfuss, 1837)

342 (1.8%), Fedaiella inaequiplicata (von Klipstein, 1843) (gastropod, 1.4%), Naticopsis seisiensis

343 (Blaschke, 1905) (gastropod 1.2%) and Heminajas fissidentata (von Wöhrmann, 1889) (1.1%; Figs 5

344 and 6). The fauna is clearly dominated by the two bivalve species Pachycardia rugosa and

345 Pachycardia plieningeri that form the trophic nucleus. All other species are relatively rare and almost

346 half of the species is only represented by one individual (Figs 8b and 11). Simpson-D ranges between

15

347 0.29 and 0.47, with an average value of 0.39 (Online Resource 1). Again, the rarefaction analysis

348 shows that the diversity of some samples is underestimated (Fig. 4b).

349 A palaeontological analysis of the fauna from the Pachycardientuffe was already performed by

350 Fürsich and Wendt (1977), based on 283 individuals belonging to 26 species, including bivalves (11

351 species), gastropods (11 species), crinoids (two species), echinoids and corals (one species,

352 respectively). This analysis shows a similar rank-abundance pattern as in our study. Pachycardia

353 rugosa is with 85.5% the most abundant species, whereas almost three-quarters of all species are

354 represented by only one individual. Because Fürsich and Wendt (1977) did not separate Pachycardia

355 rugosa and Pachycardia plieningeri, the Simpson-D in their study is relatively higher (0.71) than in our

356 study (min. D = 0.29; max. D = 0.46). Whether P. rugosa and P. plieningeri represent two different

357 species is debatable. In this study, we used the less deeply depressed lunule and the more posterior

358 position of the beaks in P. plieningeri as criteria for their separation. Additional criteria noted by

359 Broili (1903) include a more elongated outline and more elongated, but narrower anterior cardinal

360 tooth in P. plieningeri.

361 The fauna of the Pachycardientuffe is bivalve dominated in terms of species richness and total

362 number of individuals. The guild-species diversity shows that epifaunal herbivores and/or detritus-

363 feeders, epibyssate suspension-feeders and shallow infaunal suspension-feeders are the most

364 important guilds (Fig. 8b). However, most individuals belong to shallow to moderate deep infaunal

365 suspension-feeders, which is not surprising since both dominating species belong to this group.

366

367 5.2. Interpretation of the fauna from the Pachycardientuffe

368

369 Several lines of evidence indicate that the fauna of the Pachycardientuffe is allochthonous: (1) the

370 lack of fossils that were preserved in life position; (2) the lack of bivalves that are preserved with

16

371 conjoined valves; (3) frequent fragmentation of fossils; (4) the poorly sorted sediment. These

372 observations raise the question where the transported fossils originally came from.

373 In contrast to the Schlernplateau fauna, where many species occur in similar abundances, the

374 fauna from the Pachycardientuffe is clearly dominated by only two species, namely Pachycardia

375 rugosa and Pachycardia plieningeri. A comparison between the fauna of the Schlernplateau beds and

376 the Pachycardientuffe revealed that 45% of all species occur exclusively in the Pachycardientuffe and

377 38% of all species occur exclusively in the Schlernplateau beds. The remaining 17% of all species

378 occur in the Schlernplateau beds as well as in the Pachycardientuffe (Online Resource 1). Species that

379 occur in the Schlernplateau beds as well as in the Pachycardientuffe include the bivalve species

380 Myophoria kefersteini, Heminajas fissidentata, Pachycardia rugosa, Myophoriopis richthofeni and

381 Trigonodus rablensis, which belong to the most abundant species in the Schlernplateau beds as well

382 as in the Pachycardientuffe. All these species were shallow infaunal suspension-feeders that required

383 a soft substratum. However, there are large differences in the relative abundances of these species

384 between the Schlernplateau beds and the Pachycardientuffe. Myophoria kefersteini and Heminajas

385 fissidentata are the most common species in the Schlernplateau beds but only occur rarely in the

386 Pachycardientuffe. In contrast, Pachycardia rugosa is the dominant species in the Pachycardientuffe,

387 but it ranks third in the Schlernplateau beds, where it is much less common than Myophoria

388 kefersteini and Heminajas fissidentata. If the Schlernplateau beds were the only source area for the

389 fossils of the Pachycardientuffe, then Pachycardia rugosa should be less dominant. It is therefore

390 likely that the fauna of the Pachycardientuffe did not originate solely from the Schlernplateau beds.

391 Based on palaeogeographic reconstructions (Brandner 1991, p. 12, Fig. 6; own observations) and

392 ecological features of the fauna, we identified two additional possible source areas for the fauna of

393 the Pachycardientuffe. One of these additional source areas is the shallow clastic coast of a nearby

394 volcanic island and/or adjacent submarine highs, where a fauna could have flourished in

395 volcanoclastic sediments. Brandner (1991, p. 12, Fig. 6) indicated the presence of a structural high

396 with a volcanic island topography few kilometres southeast of the Frommerbach localities, which was 17

397 caused by the tectonic activity during the Late Ladinian and consisted of uplifted volcanoclastics of

398 the Wengen Fm. Thick shelled, shallow-burrowing bivalves such as Pachycardia are typical faunal

399 elements of such shallow-marine, sandy sediments (Stanley 1970). Brandner (1991, p. 12, Fig. 6)

400 indicated dipping of the palaeo-relief of the structural high towards the northwest, directly pointing

401 towards the Frommerbach localities (see also Yose 1991, p. 35). We assume that at least parts of the

402 fauna of the Pachycardientuffe were transported from this source area via submarine high-density

403 gravity flows, together with volcanogenic material (Figs 12a-c). A similar scenario was already

404 described by Fürsich and Wendt (1977), who assumed that the high number of Pachycardia indicates

405 a distinct association in a shallow-water high-energy environment, e.g. in the vicinity of volcanic

406 swells or islands, and that this Pachycardia dominated fauna together with tuffitic material and

407 faunal elements from neighbouring biotopes was transported into the basin.

408 A third possible source area is the reef environment that fringed the lagoon of the Schlernplateau

409 beds. Blaschke (1905, p. 219) reported the presence of several bivalve taxa in the Pachycardientuffe

410 that have previously been reported from the reefal facies of the “Schlerndolomit” sensu lato. These

411 bivalves include Chlamys (Granulochlamys) cf. tubulifera (Münster, 1841), Mysidioptera cf.

412 aviculaeformis Broili, 1903, Mysidioptera cf. incurvostriata (Gümbel, 1861), Mysidioptera cf. spinigera

413 Bittner, 1895, Mysidioptera acuta Broili, 1903, Mysidioptera cf. laczkói Bittner, 1901, Cassianella

414 decussata (Münster, 1841), Waagenoperna cf. planata (Broili, 1903), Pteria aff. seisiana (Broili, 1903)

415 and Badiotella sp., which were mostly epibyssate suspension-feeders that needed a hard substratum

416 for attachment. Furthermore, gastropods from the family Patellidae were reported from the

417 Pachycardientuffe (Blaschke 1905), which also live on hard substratum.

418 Fig. 13 provides a summary of our interpretation of the fauna from the Pachycardientuffe,

419 showing transportation from the following three sources: (1) the soft ground lagoonal fauna of the

420 Schlernplateu beds, (2) the hard ground reef fauna from the Schlerndolomite and (3) a Pachycardia

421 rugosa dominated fauna from the coast from a near volcanic island.

18

422 As mentioned above, the number of species found in this study (59 species) is much lower than

423 the number of species described by Broili (1903), Blaschke (1905), Waagen (1903) and Waagen

424 (1907; altogether more than 270 species). One explanation for the high number of species described

425 in these monographs is taxonomic oversplitting. A taxonomic revision of the fauna is beyond the

426 scope of this study, but an inspection of Broili’s (1903) type material in Munich confirms that his

427 taxonomic approach was more typological than usually applied today, and that e.g. the species

428 richness of the limid bivalve genus Mysidioptera (24 reported species) has been clearly

429 overestimated. However, also the number of reported genera is higher in the literature than in our

430 samples (100 versus 38 genera), thus oversplitting of species is not the sole reason for the difference

431 in reported diversity. An additional factor that contributed to the higher species richness in the

432 historical monographs is probably that these were based on a higher number of geographically

433 distant sampling localities, which likely had different source areas and thus differed considerably in

434 their taxonomic composition. Therefore, the lower species richness in our samples might by largely

435 due to the poorer present-day outcrop conditions, which limited our field collections to two nearby

436 localities.

437

438 6. Summarized interpretation of the benthic faunas from the Schlernplateau beds and

439 Pachycardientuffe

440

441 Our field study led to the recognition of 52 species in the Schlernplateau beds and 59 species in

442 the Pachycardientuffe. The sedimentological and palaeontological analyses outlined above allow the

443 detailed palaeoecological interpretation of these faunas.

444 The recorded species from the Schlernplateau beds lived in a lagoon, close to an oceanic inlet. The

445 fauna is unaffected from mixing by transportation, but it was probably subject to some time-

19

446 averaging. This means that some of the recorded species did not live contemporaneously at the same

447 place, thus the overall number of species is a certain overestimation of the community species

448 richness at a given time. However, the overall diversity of the Schlernplateau fauna represents a

449 good estimation of the taxonomic richness of shelly soft-bottom dwellers in a shallow-water setting

450 of a tropical sea at the end of the Middle Triassic.

451 In contrast to the Schlernplateau beds, the fauna of the Pachycardientuffe represents almost

452 certainly palaeocommunities from at least three different settings (lagoon, reef and sandy shallow

453 sea) that were mixed during transportation into the basin. The fact that our collection contains

454 significantly less species (59) than recorded in the literature (>270) is largely due to the lower

455 number of sampled localities that encompass a lower number of ecologically different source areas.

456 The evidence for faunal mixing in the Pachycardientuffe does not invalidate the data for studies of

457 biodiversity during the Middle Triassic, but it poses some constraints on their interpretation. The

458 fauna of the Pachycardientuffe does not represent a single palaeocommunity, but an integral of the

459 shallow marine shelly fauna that lived in different habitats at the same time. It is thus a good

460 estimator for the gamma-diversity of the reef-to-basin landscape of the tropical Tethyan ocean

461 during the late Middle Triassic.

462

463 7. Species richness of Early and Middle Triassic benthic faunas: a comparison

464

465 To what extend do Early and Middle Triassic benthic communities differ in their ecology, species

466 richness and taxonomic composition? Answering this question is crucial for identifying evolutionary

467 processes during the main phase of rediversification after the end-Permian mass extinction. We here

468 present a preliminary comparison between selected benthic communities of these two epochs. Data

469 used for this comparison are shown in Tab. 1. Lower and Middle Triassic macroinvertebrate faunas

20

470 selected for the comparison come from lithological units that represent similar shallow-marine

471 environments. Because most of the data used for this comparison are raw data from the literature,

472 synonyms and taxonomic oversplitting (see also chapter 5.2) may cause a certain bias towards

473 greater richness, especially in Middle Triassic faunas where modern revisions mostly lack. A

474 reevaluation of bivalve species from the Jena Fm. and the Meissner Fm. (Hautmann, unpublished

475 data) suggests oversplitting by nearly 50%. However, because oversplitting is likely similar in all major

476 taxa, the gastropod/bivalve ratio discussed below is probably robust, and a pronounced Anisian rise

477 in diversity maintains even if an overestimation of diversity by factor 2 is assumed.

478 The comparison between Early and Middle Triassic benthic communities shows clearly that the

479 overall diversity increased through time; however, diversity slowly increased during the Early Triassic

480 and rapidly during the Middle Triassic (Tab. 1; Fig. 14a). This pattern is evident in the trends of the

481 most diverse taxa in Early and Middle Triassic benthic communities, i.e., bivalves, gastropods and

482 brachiopods. Our compilation shows that bivalves were clearly the most diverse taxon in Early

483 Triassic benthic communities. Although bivalves were relatively diverse immediately after the end-

484 Permian mass extinction, the net increase in species richness was small until the end of the Early

485 Triassic. The major diversification of bivalves occurred during the Anisian (Hautmann 2007; Tu et al.

486 2016). Already the Late Bithynian–Early Illyrian Jena Fm. yields a highly diverse Middle Triassic

487 bivalve fauna, surpassed by the late Anisian fauna of the Leidapo Mb. (Tab. 1). Gastropods were on

488 average the second most diverse taxon during the Early Triassic, and their increase in species

489 richness during that time was comparable to that of bivalves. During the Middle Triassic, however,

490 gastropod species richness increased more rapidly than bivalve species richness. In consequence,

491 gastropods were the most diverse group in many Middle Triassic lithological units (Tab. 1). The

492 change from the taxonomic dominance of bivalves in Early Triassic benthic communities to the

493 taxonomic dominance of gastropods during the Middle Triassic can also be seen in the

494 gastropod/bivalve-ratio that was variable but with an increasing trend through time (Fig. 14b).

495 Brachiopods were the third most diverse group in almost all Early and Middle Triassic benthic

21

496 communities. Their species richness was relatively low during the Early Triassic, and in most

497 Olenekian lithological units used for this comparison they are even not reported. However,

498 brachiopods also became more diverse during the Middle Triassic, although their increase in species

499 richness during the Middle Triassic is much slower compared to bivalves and gastropods.

500 The comparison between Early and Middle Triassic benthic communities confirms that corals

501 were absent in the entire Early Triassic (e.g., Flügel 2002; Weidlich et al. 2003; Stanley 2003; Payne et

502 al. 2006; Brayard et al. 2011). Sponges occurred only in two Lower Triassic formations used for this

503 comparison, but sponges, in association with microbialites, are known from the Lower Triassic

504 (Induan) of Armenia (Friesenbichler et al. 2018), Lower and Middle Triassic carbonate successions of

505 the Western Tethys domain (the Alps, Carpathians and the Germanic Basin; SculzSzulc 2007) as well

506 as Lower Triassic strata from China (Ezaki et al. 2008; Bagherpour et al. 2017) and the Western US

507 (Brayard et al. 2011; Hofmann et al. 2014). However, the number of these sponge species is not

508 known. Middle Triassic sponges are not only reported in the lithological units shown in Tab. 1 but

509 also in the Illyrian strata of the Wetterstein Fm. (Velledits et al. 2011). The fossil record of

510 echinoderms in the Early to Middle Triassic is heterogeneous. Several Palaeozoic clades of

511 echinoderms survived the end-Permian mass extinction and reappeared as Lazarus taxa in the

512 Middle Triassic, without giving rise to a new diversification (Thuy et al. 2017). Crinoid diversity was

513 very low during the Early Triassic (Hagdorn 2011), although cladistics analyses suggest that initial

514 diversification took place in the Induan (Twitchett and Oji 2005) or already in the Late Permian (Oji

515 and Twitchett (2015); however, well before the documented species richness increase during the

516 Middle and early Late Triassic (Hagdorn 2011). Only two echinoid genera are reported from Lower

517 Triassic strata (Twitchett and Oji 2005) and very low levels of diversity were apparently maintained

518 for at least the duration of the Early and Middle Triassic. Nevertheless, numerous undescribed

519 echinoid spines and other remains have been noted in Early Triassic fossil assemblages (e.g., from the

520 Griesbachian of Oman; Krystyn et al. 2003) and these undescribed records may hint at a hidden

521 diversity of echinoids within the Early Triassic (see also Thompson et al. 2015; Thuy et al. 2017). Fossil

22

522 remains or trace fossils attributed to Lower Triassic asteroids have only been recorded from

523 Olenekian strata of the western USA (Schubert and Bottjer 1995; Villier et al. 2018). Definite reports

524 from Lower Triassic holothuroids only exist from Western Nepal (Kozur and Mostler 1973; Mostler

525 1977), whereas a total number of 84 holothuroid species was reported from several Middle Triassic

526 strata (Reich 2013). Fossil remains from ophiuroids are quite abundant in Lower Triassic strata

527 worldwide but so far only six species have been described (Twitchett and Oji 2005). Early Triassic

528 bryozoans were reported from a couple of Lower Triassic strata (Nakrem and Mørk 1991; Wignall and

529 Twitchett 2002; Baud et al. 2008) but only five Early Triassic bryozoan genera are reported (Powers

530 and Pachut 2007) and only six species have been described formally (Baud et al. 2008); however,

531 bryozoan diversity increased during the Middle and Late Triassic, with eight reported genera (Powers

532 and Pachut 2007) a total number of more than 40 species (see reviews in Schäfer and Grant-Mackie

533 1994; Schäfer et al. 2003). Microconchids were important constituents of marine ecosystems

534 immediately after the Permian/Triassic mass extinction (He et al. 2012) and were also commonly

535 present through the entire Early Triassic worldwide (Fraiser 2011; Zatoń et al. 2013b; Yang et al.

536 2015). Although microconchids can be very abundant locally, they are lowly diverse and only

537 represented by three genera (Zatoń et al. 2013b).

538 The Middle Triassic increase in species richness might be related to more stable environmental

539 conditions, as reflected by the constancy of the Middle Triassic carbon isotope curve (Payne et al.

540 2004), or alternatively represent the effect of competition-driven niche partitioning after

541 rediversification exceeded a threshold (Hautmann 2014; Hautmann et al. 2015). The change in the

542 gastropod/bivalve-ratio is more difficult to interpret, but it is notable that occurrences of high

543 gastropod species richness relative to bivalves are almost exclusively linked to carbonate platform

544 environments (Fig. 14b). This might represent a taphonomic bias due to differences in skeletal

545 mineralogy between gastropods and bivalves, but we note that aragonite-shelled bivalves are

546 commonly preserved in the Early Triassic, which makes this explanation unlikely. Alternatively, we

547 hypothesise that the increase in gastropod species richness is a primary signal linked to the

23

548 resurgence of large carbonate platform and reefs during the Middle Triassic (Gaetani et al. 1981;

549 Senowbari et al. 1993), although details of the ecological causes for this facies-related increase in

550 gastropod species richness requires further investigation.

551

552 8. Conclusion

553

554 Early and Middle Triassic shallow-marine benthic communities differed considerably in terms of

555 species richness and taxonomic composition. Whereas Early Triassic formations from shallow-marine

556 settings contain generally less than 50 macroinvertebrate species, regional diversities of around 300

557 species have been described from various Anisian-Ladinian shallow-marine settings worldwide (Tab.

558 1; Fig. 14), including the herein studied fauna from the Pachycardientuffe. For more narrowly defined

559 habitats, such as that of the open lagoon represented by the Schlernplateau fauna (85 species), the

560 increase in diversity was likewise remarkable. The Middle Triassic increase in diversity at the

561 local/regional level analysed herein correlates with the most pronounced increase in bivalve diversity

562 during the entire Triassic in a global dataset (Hautmann 2007, fig. 8) and therefore represents the

563 main phase of recovery after the end-Permian mass extinction. This main phase of recovery in

564 benthic marine communities was associated with a weakening of the dominance of bivalves in

565 comparison to the Early Triassic, which reflects the proliferation of other invertebrate taxa,

566 particularly gastropods. We hypothesise that the resurgence of carbonate platforms during the

567 Middle Triassic played an important role in the shift of taxonomic compositions, and that the

568 increase of environmental diversity associated with newly arising carbonate platforms contributed at

569 least partly to the accelerated pace of recovery during the Middle Triassic.

24

570 References

571 Formatted: Justified, Indent: Left: 0", First line: 0"

572 Aberhan, M. 1994. Guild-Structure and evolution of Mesozoic Benthic shelf Communities. Palaios 9:

573 516-–545.

574 Alroy, J., M. Aberhan, D.J. Bottjer, M. Foote, F.T. Fürsich, P.J. Harries, A.J.W. Hendy, S.M. Holland, L.C.

575 Ivany, W. Kiessling, M.A. Kosnik, C.R. Marshall, A.J. McGowan, A.I. Miller, T.D. Olszewski, M.E.

576 Patzkowsky, S.E. Peters, L. Villier, P.J. Wagner, N. Bonuso, P.S. Borkow, B. Brenneis, M.E. Clapham,

577 L.M. Fall, C.A. Ferguson, V.L. Hanson, A.Z. Krug, K.M. Layou, E.H. Leckey, S. Nürnberg, C.M.

578 Powers, J.A. Sessa, C. Simpson, A. Tomašových, and C.C. Visaggi. 2008. Phanerozoic trends in the

579 global diversity of marine invertebrates. Science 321: 97-–100.

580 Autonome Provinz Bozen (ed.). 2007. Geologische Karte der Westlichen Dolomiten.

581 Bagherpour, B., H. Bucher, A. Baud, M. Brosse, T. Vennemann, R. Martin, and K. Guodun. 2017.

582 Onset, development, and cessation of basal Early Triassic microbialites (BETM) in the Nanpanjiang

583 pull-apart Basin, South China Block. Gondwana Research 44: 178-–204.

584 Balini, M., S.G. Lucas, J.F. Jenks, and J.A. Spielmann. 2010. Triassic ammonoid biostratigraphy: an

585 overview. In The Triassic Timescale, ed. S.G. Lucas, 221-–262. Geological Society of London, Special

586 Publications 334.

587 Batten, R.L., and W.L. Stokes, W.L. 1986. Early Triassic gastropods from the Sinbad Member of the

588 Moenkopi Formation, San Rafael Swell, Utah. American Museum Novitates 2864: 1-–33.

589 Baud, A., H.A. Nakrem, B. Beauchamp, T.W. Beatty, A.F. Embry, and C.M. Henderson. 2008. Lower

590 Triassic bryozoan beds from Ellesmere Island, High Arctic, Canada. Polar Research 27: 428-–440.

591 Bittner, A. 1890. Brachiopoden der alpinen Trias. Abhandlungen der k. k. geologischen Reichsanstalt

592 14: 1-–325.

25

593 Bittner, A. 1892. Brachiopoden der alpinen Trias - Nachtrag. Abhandlungen der k. k. geologischen

594 Reichsanstalt 17: 1-–55.

595 Bittner, A. 1895. Lamellibranchiaten der alpinen Trias. I Theil: Revision der Lamellibranchiaten von

596 Sct. Cassian. Abhandlungen der k. k. Geologischen Reichsanstalt 18: 1-–236.

597 Bittner, A. 1900. Ueber die triadische Lamellibranchiaten-Gattung Mysidioptera Sal. und deren

598 Beziehungen zu palaeozoischen Gattungen. Jahrbuch der k. k. Geologischen Reichsanstalt 50: 59-–

599 66.

600 Bittner, A. 1901. Lamellibranchiaten aus der Trias des Bakonyer Waldes. Resultate der

601 wissenschaftlichen Erforschung des Balatonsees, Anhang: Palaeontologie der Umgebung des

602 Balatonsees II, Band 3: 1-–107.

603 Blaschke, F. 1905. Die Gastropodenfauna der Pachycardientuffe der Seiseralpe in Südtirol nebst

604 einem Nachtrag zur Gastropodenfauna der roten Raibler Schichten auf dem Schlernplateau.

605 Beiträge zur Paläontologie und Geologie Österreich-Ungarns und des Orients 17: 161-–221.

606 Böhm, J. 1895. Die Gastropoden des Marmolatakalkes. Palaeontographica 42: 211-–308.

607 Bosellini, A. 1998. Geologie der Dolomiten. Bozen: Verlagsanstalt Athesia.

608 Bottoli, G., and S. Trombetta. 1998. Analisi di facies ed ambiente deposizionale del Conglomerato Formatted: English (United States)

609 della Marmolada: un sistema torbiditico dominato da flussi ad alta densità (gruppo Col Rossi-Porta

610 Vescovo, Dolomti occidentali). Memorie Società Geologica Italiana 53: 341-–357. Field Code Changed

611 Brack, P., and H. Rieber. 1993. Toward a better definition of the Anisian/Ladinian boundary: New

612 biostratigraphic data and correlations of boundary sections from the Southern Alps. Eclogae

613 Geologicae Helvetiae 86: 415-–527.

26

614 Brack, P., R. Mundil, F. Oberli, M. Meier, and H. Rieber. 1996. Biostratigraphic and radiometric age

615 data question the Milankovitch characteristics of the Latemar cycles (Southern Alps, Italy).

616 Geology 24: 371-–375.

617 Brandner, R. 1991. Geological Setting and Stratigraphy of the Schlern-Rosengarten Buildup and Seiser

618 Alm Basin. In The Northern Margin of the Schlern/Sciliar-Rosengarten/Catinaccio Platform, eds. R.

619 Brandner, E. Flügel, R. Koch, and L.A. Yose, 4-–16. Dolomieu Conference on Carbonate Platforms

620 and Dolomitization, Guidebook Excursion A, Ortisei (Italy).

621 Brandner, R., A. Gruber, and L. Keim. 2007. Geologie der westlichen Dolomiten: Von der Geburt der

622 Neotethys im Perm zu Karbonatplattformen, Becken und Vulkaniten der Trias. Geo.Alp 4: 95-–121.

623 Brandner, R., A. Gruber, C. Morelli, and V. Mair. 2016. Pulses of Neotethys-Rifting in the

624 Permomesozoic of the Dolomites. Geo.Alp 13: 7-–70.

625 Brayard, A., E. Vennin, N. Olivier, K G. Bylund, J. Jenks, D.A. Stephen, H. Bucher, R. Hofmann, N.

626 Goudemand, and G. Escarguel. 2011. Transient metazoan reefs in the aftermath of the end-

627 Permian mass extinction. Nature Geoscience 4: 693-–697.

628 Brayard, A., G. Escarguel, H. Bucher, C. Monnet, T. Brühwiler, N. Goudemand, T. Galfetti, and J. Guex.

629 2009. Good genes and good luck: ammonoid diversity and the end-Permian mass extinction.

630 Science 325: 1118-–1121.

631 Brayard, A., L.J. Krumenacker, J.P. Botting, J.F. Jenks, K.G. Bylund, E. Fara, E. Vennin, N. Olivier, N.

632 Goudemand, T. Saucède, S. Charbonnier, C. Romano, L. Doguzhaeva, B. Thuy, M. Hautmann, D.A.

633 Stephen, C. Thomazo, G. and Excarguel. 2017. Unexpected Early Triassic marine ecosystem and

634 the rise of the Modern fauna. Science Advances 3: e1602159.

635 Broglio Loriga, C., C. Neri, and R. Posenato. 1986. The Early Macrofaunas of the Werfen Formation

636 and the Permian-Triassic Boundary in the Dolomites (Southern Alps, Italy). Acta Geologica 62: 3-–

637 18. 27

638 Broglio Loriga, C., F. Góczán, J. Haas, K. Lenner, C. Neri, A. Oravecz-Scheffer, R. Posenato, I. Szabó,

639 and Á. Tóth-Makk. 1990. The Lower Triassic sequences of the Dolomites (Italy) and Transdanubian

640 Mid-Mountains (Hungary) and their correlation. Memorie di Scienze Geologiche 42: 41-–103.

641 Broili, F. 1903. Die Fauna der Pachycardientuffe der Seiser Alp (Mit Ausschluss der Gastropoden und

642 Cephalopoden). Palaeontographica 50: 145-–227.

643 Budai, T. 2008. Geological excursions in the Alpine and Germanic Triassic facies areas of Hungary.

644 Triassic Workshop 2008, Hungary: 1-–50.

645 Busse, E. 1972. Fazies und Fauna des Oberen Muschelkalkes von Willebadessen. Philippia 1: 110-–

646 126.

647 Eichwald, ME. 1851. Naturhistorische Bemerkungen als Beitrag zur vergleichenden Geognosie, auf

648 einer Reise durch die Eife, TirolEifel, Tyrol, Italien, Sizilien und Algier. II. Geognostischer Ausflug

649 nach Tyrol. Nouveaux mémoires de la Société impériale des naturalistes de Moscou 9: 75-–204.

650 Ezaki Y., J. Liu, T. Nagano and N. Adachi. 2008. Geobiological aspects of the earliest Triassic

651 microbialites along the southern periphery of the tropical Yangtze Platform: initiation and

652 cessation of a microbial regime. Palaios 23: 356-–369.

653 Flügel, E. 2002. Triassic reef patterns. In Phanerozoic Reef Patterns, eds. W. Kiessling, E. Flügel, and J.

654 Golonka, 391-–463. SEPM Special Publication 72, Society for Sedimentary Geology.

655 Foster, W.J., and K. Sebe. 2017. Recovery and diversification of marine communities following the

656 late Permian mass extinction event in the western Palaeotethys. Global and Planetary Change

657 155: 165-–177.

658 Foster, W.J., S. Danise, A. Sedlacek, G.D. Price, K. Hips, and R.J. Twitchett. 2015. Environmental

659 control on the post-Permian recovery of benthic, tropical marine ecosystems in western

28

660 Palaeotethys (Aggtelek Karst, Hungary). Palaeogeography, Palaeoclimatology, Palaeoecology 440:

661 374-–394.

662 Foster, W.J., S. Danise, G.D. Price, and R.J. Twitchett. 2017. Subsequent biotic crises delayed marine

663 recovery following the late Permian mass extinction event in northern Italy. PLoS One 12:

664 e0172321.

665 Fraiser, M.L. 2011. Paleoecology of second tiers from western Pangean tropical marine environments

666 during the aftermath of the end-Permian mass extinction. Palaeogeography, Palaeoclimatology,

667 Palaeoecology 308: 181-–189.

668 Friesenbichler, E., S. Richoz, A. Baud, L. Krystyn, L. Sahakyan, S. Vardanyan, J. Peckmann, J. Reitner,

669 and K. Heindel. 2018. Sponge-microbial build-ups from the lowermost Triassic Chanakhchi section

670 in southern Armenia: Microfacies and stable carbon isotopes. Palaeogeography,

671 Palaeoclimatology, Palaeoecology 490: 653-–672.

672 Frizzell, D.L., and H. Exline. 1956. Monograph of Fossil Holothurian Sclerites. Bulletin of School of

673 Mines and Metallurgy (Technical Series) 89: 1-–204.

674 Fürsich, F.T. 1994. Palaeoecology and evolution of Mesozoic salinity-controlled benthic

675 macroinvertebrate associations. Lethaia 26: 327-–346.

676 Fürsich, F.T., and J. Wendt. 1977. Biostratinomy and palaeoecology of the Cassian Formation

677 (Triassic) of the Southern Alps. Palaeogeography, Palaeoclimatology, Palaeoecology 22: 257-–323.

678 Fürsich, F.T., and M. Aberhan. 1990. Significance of time‐averaging for palaeocommunity analysis.

679 Lethaia 23: 143-–152.

680 Fürsich, F.T., S. Freytag, J. Röhl, and A. Schmid. 1995. Palaeoecology of benthic associations in

681 salinity-controlled marginal marine environments: Examples from the Lower Bathonian (Jurassic)

29

682 of the Causses (southern France). Palaeogeography, Palaeoclimatology, Palaeoecology 113: 135-–

683 172.

684 Gaetani, M., E. Fois, F. Jadoul, and A. Nicora. 1981. Nature and evolution of Middle Triassic carbonate

685 buildups in the Dolomites (Italy). Marine Geology 44: 25-–57.

686 Goldfuss, G.A. 1833-1841. Petrefacta Germaniae. Teil 2. 1833: pp. 1-–68, pls. 72-–96; 1835: pp. 69-–

687 140, pls. 97-–121; 1837: pp. 141-–224, pls. 122-–146; 1841: pp. 225-–312, pls. 147-–165;

688 Düsseldorf (Arnz & Co).

689 Gredler, V.M. 1862. Vierzehn Tage in Bad Ratzes: eine naturgeschichtliche Lokalskizze mit näherer

690 Berücksichtigung der Fauna. Programm des Kaiserlich-Königlichen Gymnasiums zu Bozen,

691 veröffentlicht am Schlusse des Schuljahres 13 (1862/63): 1-–41.

692 Gümbel, C.W. von. 1861. Geognostische Beschreibung des bayerischen Alpengebirges und seines

693 Vorlandes. Gotha: Verlag von Justus Perthes.

694 Häberle, D. 1908. Paläontologische Untersuchung triadischer Gastropoden aus dem Gebiet von

695 Predazzo. Verhandlungen des Naturhistorisch-Medizinischen Vereins zu Heidelberg 9: 1-–630.

696 Hagdorn, H. 2011. Triassic: the crucial period of post-Palaeozoic crinoid diversification. Swiss Journal

697 of Palaeontology 130: 91-–122.

698 Hagdorn, H., and E. Nitsch. 2009. 6th International Triassic Field Workshop (Pan-European Correlation

699 of the Triassic). Triassic of Southwest Germany. Freiburg: Landesamt für Geologie, Rohstoffe und

700 Bergbau.

701 Hagdorn, H., and R. Mundlos. 1982. Autochthonschille im Oberen Muschelkalk (Mitteltrias)

702 Südwestdeutschlands. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 162: 332-–

703 351.

30

704 Hagdorn, H., and W. Ockert. 1993. Encrinus liliiformis im Trochitenkalk Süddeutschlands. In:

705 Muschelkalk. Sonderbände der Gesellschaft für Naturkunde in Württemberg e.V. 2, 245-–260.

706 Stuttgart, Korb: Goldschneck-Verlag Werner K. Weidert.

707 Hammer, Ø., D.A.T. Harper, and P.D. Ryan. 2001. PAST: Paleontological Statistics Software Package

708 for Education and Data Analysis. Palaeontologia Electronica 4: 1-–9.

709 Hauer, F. von. 1857. Ein Beitrag zur Kenntniss der Fauna der Raibler Schichten. Sitzungsberichte der

710 Kaiserlichen Akademie der Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klasse

711 24: 1-–32.

712 Hausmann, I.M., A. and Nützel. 2015. Diversity and palaeoecology of a highly diverse Late Triassic

713 marine biota from the Cassian Formation of north Italy. Lethaia 48: 235-–255.

714 Hautmann, M. 2007. Steuerungsfaktoren globaler Biodiversifizierungsmuster am Beispiel

715 frühmesozoischer Muscheln. Beringeria 37: 61-–74.

716 Hautmann, M. 2014. Diversification and diversity partitioning. Paleobiology 40: 162-–176.

717 Hautmann, M., A.B. Smith, A.J. McGowan, and H. Bucher. 2013. Bivalves from the Olenekian (Early

718 Triassic of south-western Utah: systematics and evolutionary significance. Journal of Systematic

719 Palaeontology 11: 263-–293.

720 Hautmann, M., and H. Hagdorn. 2013. Oysters and oyster-like bivalves from the Middle Triassic

721 Muschelkalk of the Germanic Basin. Paläontologische Zeitschrift 87: 19-–32.

722 Hautmann, M., B. Bagherpour, M. Brosse, Å. Frisk, R. Hofmann, A. Baud, A. Nützel, N. Goudemand,

723 and H. Bucher. 2015. Competition in slow motion: The unusual case of benthic marine

724 communities in the wake of the end-Permian mass extinction. Palaeontology 58: 871-–901.

31

725 Hautmann, M., H. Bucher, T. Brühwiler, N. Goudemand, A. Kaim, and A. Nützel. 2011. An unusually

726 diverse mollusc fauna from the earliest Triassic of South China and its implications for benthic

727 recovery after the end-Permian biotic crisis. Geobios 44: 71-–85.

728 He, L., Y.B. Wang, A. Woods, G.S. Li, H. Yang, and W. Liao. 2012. Calcareous tubeworms as disaster

729 forms after the end-Permian mass extinction in South China. Palaios 27: 878–886.

730 Hofmann, R., M. Hautmann, A. Brayard, A. Nützel, K.G. Bylund, J.F. Jenks, E. Vennin, N. Olivier, and H.

731 Bucher. 2014. Recovery of benthic marine communities from the end‐Permian mass extinction at

732 the low latitudes of eastern Panthalassa. Palaeontology 57: 547-–589.

733 Hofmann, R., M. Hautmann, and H. Bucher. 2013a. A new paleoecological look at the Dinwoody

734 Formation (Lower Triassic, Western USA): Intrinsic versus extrinsic controls on ecosystem

735 recovery after the end-Permian mass extinction. Journal of Paleontology 87: 854-–880.

736 Hofmann, R., M. Hautmann, and H. Bucher. 2015. Recovery dynamics of benthic marine communities

737 from the Lower Triassic Werfen Formation, northern Italy. Lethaia 48: 474-–496.

738 Hofmann, R., M. Hautmann, M. Wasmer, and H. Bucher. 2013b. Palaeoecology of the Spathian Virgin

739 Formation (Utah, USA) and its implications for the Early Triassic recovery. Acta Palaeontologica

740 Polonica 58: 149-–173.

741 Keim, L. 2008. Geologie im Gebiet Schlern-Seiser Alm: vom Tethysmeer zum Gebirge. Gredleriana 8:

742 25-–46.

743 Kittl, E. 1891. Die Gastropoden der Schichten von St. Cassian der südalpinen Trias. I. Theil. Annalen

744 des Naturhistorischen Museums in Wien 6: 166-–262.

745 Kittl, E. 1892. Die Gastropoden der Schichten von St. Cassian der südalpinen Trias. II. Theil. Annalen

746 des Naturhistorischen Museums in Wien 7: 35-–97.

32

747 Kittl, E. 1894. Die Gastropoden der Schichten von St. Cassian der südalpinen Trias. III. Theil. Annalen

748 des Naturhistorischen Museums in Wien 9: 143-–277.

749 Klipstein, A. von 1843-1845. Mittheilungen aus dem Gebiete der Geologie und Palaeontologie. 1843:

750 pp. 1-–144, pls. 1-–9; 1844: pp. 145-–240, pls. 10-–15; 1845: pp. 241-–311, pls. 16-–20; Gießen

751 (Heyer). 3. Abt. (1883): 1-–101, 3 Tafeln; Gießen (Ricker).

752 Komatsu, T., J.H. Chen, M.Z. Cao, F. Stiller, and H. Naruse. 2004. Middle Triassic (Anisian) diversified

753 bivalves: depositional environments and bivalve assemblages in the Leidapo Member of the

754 Qingyan Formation, southern China. Palaeogeography, Palaeoclimatology, Palaeoecology 208:

755 207-–223.

756 Kozur, H., and H. Mostler. 1973. Beiträge zur Mikrofauna permotriadischer Schichtfolgen Teil 1:

757 Conodonten aus der Tibetzone des Niederen Himalaya (Dolpogebiet, Westnepal). Geologische

758 und Paläontologische Mitteilungen Innsbruck 3: 1–23.

759 Krystyn, L., A. Baud, S. Richoz, and R.J. Twitchett. 2003. A unique Permian-Triassic boundary section

760 from Oman. Palaeogeography, Palaeoclimatology, Palaeoecology 191: 329-–344.

761 Laube, G.C. 1865a. Die Fauna der Schichten von St. Cassian: Ein Beitrag zur Paläontologie der alpinen

762 Trias, I. Abtheilung, Spongitarien, Corallen, Echiniden und Crinoiden. Wien: Kaiserlich-königliche

763 Hof- und Staatsdruckerei.

764 Laube, G.C. 1865b. Die Fauna der Schichten von St. Cassian: Ein Beitrag zur Paläontologie der alpinen

765 Trias, II. Abtheilung, Brachiopoden und Bivalven. Wien: Kaiserlich-königliche Hof- und

766 Staatsdruckerei.

767 Laube, G.C. 1868. Die Fauna der Schichten von St. Cassian: Ein Beitrag zur Paläontologie der alpinen

768 Trias, III. Abtheilung, Gastropoden – I. Hälfte. Wien: Kaiserlich-königliche Hof- und

769 Staatsdruckerei.

33

770 Laube, G.C. 1869a. Die Fauna der Schichten von St. Cassian: Ein Beitrag zur Paläontologie der alpinen

771 Trias, IV. Abtheilung, Gastropoden – II. Hälfte. Wien: Kaiserlich-königliche Hof- und

772 Staatsdruckerei.

773 Laube, G.C. 1869b. Die Fauna der Schichten von St. Cassian: Ein Beitrag zur Paläontologie der alpinen

774 Trias, V. Abtheilung, Cephalopoden. – Schluss. Wien: Kaiserlich-königliche Hof- und

775 Staatsdruckerei.

776 Leonardi, P., and F. Fiscon. 1947. Nuove forme di gasteropodi della fauna cassiana di Cortina

777 d’Ampezzo. Memorie del Museo di Storia Naturale della Venezia Tridentina 8: 1-–16.

778 Leonardi, P., and F. Fiscon. 1959. La fauna cassiana di Cortina d’Ampezzo. Parte III: Gasteropodi.

779 Memorie degli Istituti di Geologia e Mineralogia dell’Università di Padova 21: 1-–140. Formatted: German (Switzerland) Formatted: German (Switzerland)

780 Manfrin, S., P. Mietto, and N. Preto. 2005. Ammonoid biostratigraphy of the Middle Triassic Latemar Formatted: English (United States)

781 platform (Dolomites, Italy) and its correlation with Nevada and Canada. Geobios 38: 477-–504.

782 McGowan, A.J., A.B. Smith, and P.D. Taylor. 2009. Faunal diversity, heterogeneity and body size in the

783 Early Triassic: testing post-extinction paradigms in the Virgin Limestone of Utah, USA. Australian

784 Journal of Earth Sciences 56: 859-–872.

785 Morisita, M. 1959. Measuring interspecific association and similarity between communities.

786 Memoires of the faculty of Science Kyushu University series E (Biology) 3: 65-–80.

787 Mostler, H. 1977. Zur Palökologie triadischer Holothurien (Echinodermata). Berichte des

788 naturwissenschaftlich-medizinischen Vereins in Innsbruck 64: 13–40.

789 Münster, G. 1841. Beiträge zur Geognosie und Petrefacten-Kunde des südöstlichen Tirols vorzüglich

790 der Schichten von St. Cassian. II. Beschreibung und Abbildung der in den Kalkmergelschichten von

791 St. Cassian gefunden Versteinerungen. Beiträge zur Petrefacten-Kunde 4: 25-–152.

34

792 Nakrem, H.A., and A. Mørk. 1991. New Early Triassic Bryozoa (Trepostomata) from Spitsbergen, with Formatted: English (United States)

793 some remarks on the stratigraphy of the investigated horizons. Geological Magazine 128: 129-–

794 140.

795 Neri, C., and R. Posenato. 1985. New biostratigraphical data on uppermost Werfen Formation of

796 western Dolomites (Trento, Italy). Geologisch-Paläontologische Mitteilungen Innsbruck 14: 83-–

797 107.

798 Neyman, A.A. 1967. Limits to the application of the trophic group concept in benthic studies.

799 Oceanology 7: 149-–155.

800 Nützel, A., and A. Kaim. 2014. Diversity, palaeoecology and systematics of a marine fossil assemblage

801 from the Late Triassic Cassian Formation at Settsass Scharte, N Italy. Paläontologische Zeitschrift

802 88: 405-–431.

803 Nützel, A., and C. Schulbert. 2005. Facies of two important Early Triassic gastropod lagerstätten:

804 implications for diversity patterns in the aftermath of the end-Permian mass extinction. Facies 51:

805 480-–500.

806 Ockert, W. 1993. Die Zwergfaunaschichten (Unterer Hauptmuschelkalk, Trochitenkalk, mo1) im

807 nordöstlichen Baden-Württemberg. In: Muschelkalk. Sonderbände der Gesellschaft für

808 Naturkunde in Württemberg e.V. 2, 117-–215. Stuttgart, Korb: Goldschneck-Verlag Werner K.

809 Weidert.

810 Oji, T., and J. Twitchett. 2015. The oldest Post-Palaeozoic Crinoid and Permian-Triassic Origins of the Formatted: English (United States)

811 Articulata (Echinodermata). Zoological Science 32: 211-–215.

812 Payne, J. 2005. Evolutionary dynamics of gastropod size across the end-Permian extinction and

813 through the Triassic recovery interval. Paleobiology 31: 269-–290.

35

814 Payne, J.L., and M.E. Clapham. 2012. End-Permian mass extinction in the oceans: an ancient analog

815 for the twenty-first century? Annual Review of Earth and Planetary Sciences 40: 89-–111.

816 Payne, J.L., D.J. Lehrmann, J. Wei, M.J. Orchard, D.P. Schrag, and A.H. Knoll. 2004. Large

817 perturbations of the carbon cycle during recovery from the end-Permian extinction. Science 305:

818 506-–509.

819 Payne, J.L., D.J. Lehrmann, S. Christensen, J. Wei, and A.H. Knoll. 2006. Environmental and biological

820 controls on the initiation and growth of a Middle Triassic (Anisian) reef complex on the Great Bank

821 of Guizhou, Guizhou Province, China. Palaios 21: 325-–343.

822 Pia, J. 1937. Stratigraphie und Tektonik der Pragser Dolomiten in Südtirol. Wien: Eigenverlag.

823 Powers, C., and J.F. Pachut. 2008. Diversity and distribution of Triassic bryozoans in the aftermath of

824 the end-Permian mass extinction. Journal of Paleontology 82: 362-–371.

825 Raup, D.M. 1979. Size of the Permo-Triassic bottleneck and its evolutionary implications. Science 206:

826 217-–218.

827 Reich, M. 2013. How many species of fossil holothurians are there? In Echinoderms in a Changing

828 World, ed. C. Johnson, 23–51. Proceedings of the 13th International Echinoderm Conference,

829 University of Tasmania, Hobart Tasmania, Australia.

830 Richthofen, F. von. 1860. Geognostische Beschreibung der Umgegend von Predazzo, Sanct Cassian

831 und der Seiser Alpe in Süd-Tyrol. Gotha: Perthes.

832 Salomon, W. 1895. Geologische und palaeontologische Studien über die Marmolata.

833 Palaeontographica 42: 1-–210.

834 Sano, H., and K. Nakashima. 1997. Lower Triassic (Griesbachian) microbial bindstone-cementstone

835 facies, southwest Japan. Facies 36: 1-24.

36

836 Schäfer, P., and J. Grant-Mackie. 1994. Triassic Bryozoa from the Murihiku and Torlesse Supergroups,

837 New Zealand. Memoirs of the Association of Australasian Palaeontologists 16: 1-–52.

838 Schäfer, P., R.J. Cuffey, and A.R. Young. 2003. New trepostome Bryozoa from the Early Triassic

839 (Smithian/Spathian) of Nevada. Paläontologische Zeitschrift 77: 323-–340.

840 Schmidt, M. 1928. Die Lebewelt unserer Trias. Öhringen: Hohenlohesche Buchhandlung F. Rau.

841 Schmidt, M. 1938. Die Lebewelt unserer Trias - Nachtrag. Öhringen: Hohenlohesche Buchhandlung F.

842 Rau.

843 Schubert, J.K., and D.J. Bottjer. 1995. Aftermath of the Permian-Triassic mass extinction event:

844 Paleoecology of Lower Triassic carbonates in the western USA. Palaeogeography,

845 Palaeoclimatology, Palaeoecology 116: 1-–39.

846 Senowbari-Daryan, B., R. Zühlke, T. Bechstädt, and E. Flügel. 1993. Anisian (Middle Triassic) Buildups

847 of the Northern Dolomites (Italy): The Recovery of Reef Communities after the Permain/Triassic

848 Crisis. Facies 28: 181-–256.

849 Sepkoski, J.J. 1997. Biodiversity: past, present, and future. Journal of Paleontology 71: 533-–539.

850 Shigeta, Y., J.D. Zakharov, H. Maeda, A.M. Popov. 2009. The Lower Triassic System in the Abrek Bay

851 Area, South Primorye, Russia. Tokyo: National Museum of Nature and Science.

852 Song, H., P.B. Wignall, Z.Q. Chen, J. Tong, D.P.G. Bond, X. Lai, X. Zhao, H. Jiang, C. Yan, Z. Niu, J. Chen,

853 H. Yang, Y. Wang. 2011. Recovery tempo and pattern of marine ecosystems after the end-Permian

854 mass extinction. Geology 39: 739-–742.

855 Stanley, G.D. 2003. The evolution of modern corals and their early history. Earth-Science Reviews 60:

856 195-225.

857 Stanley, S.M. 1970. Relation of shell form to life habits of the Bivalvia (Mollusca). Geological Society

858 of America, Memoir 125: 1-–296. 37

859 Stanley, S.M. 2016. Estimates of the magnitudes of major marine mass extinctions in earth history.

860 Proceedings of the National Academy of Sciences 113: E6325-–E6334.

861 Stiller, F. 2001. Fossilvergesellschaftungen, Paläökologie und paläosynökologische Entwicklung im

862 Oberen Anis (Mittlere Trias) von Qingyan, insbesondere Bangtoupo, Provinz Guizhou,

863 Südwestchina. Münstersche Forschung zur Geologie und Paläontologie 92: 1-–523.

864 Stiller, F., and H. Bucher. 2008. Anisian ammonoids from Qingyan, southwestern China:

865 biostratigraphical implications for the age of the Qingyan Formation. Swiss Journal of Geosciences

866 101: 547-–562.

867 Stur, D.R.J. 1868. Eine Excursion in die Umgegend von St. Cassian. Jahrbuch der Geologischen

868 Bundesanstalt 18: 529-–568.

869 Szabó, I., S. Kovács, G. Lelkes, and A. Oravecz-Scheffer. 1979. Stratigraphic investigation of a (triassic)

870 Pelsonian-Fassanian section at Felsőörs (Balaton Highland, Hungary). Rivista Italiana di

871 Paleontologia e Stratigrafia 85: 789-–806.

872 Szulc J. 2007. Sponge-microbial stromatolites and coral-sponge reef recovery in the Triassic of the

873 western Tethys domain. In The Global Triassic, eds. S.G. Lucas and J.A. Spielmann, 402. New

874 Mexico Museum of Natural History and Science Bulletin 41.

875 Thompson, J.R., E. Petsios, E.H. Davidson, E.M. Erkenbrack, F. Gao, and D.J. Bottjer. 2015.

876 Reorganisation of sea urchin gene regulatory networks at least 268 million years ago as revealed

877 by oldest fossil cidaroid echinoid. Scientific Reports 5: 15541.

878 Thuy, B., H. Hagdorn, and A.S. Gale. 2017. Paleozoic echinoderm hangovers: Waking up in the

879 Triassic. Geology 45: 531-–534.

38

880 Tu, C., Z.Q. Chen, and D.A.T. Harper. 2016. Permian-Triassic evolution of the Bivalvia: Extinction-

881 recovery patterns linkes to ecological and taxonomic selectivity. Palaeogeography,

882 Palaeoclimatology, Palaeoecology 459: 53-–62.

883 Twitchett, R.J., and P.B. Wignall. 1996. Trace fossils and the aftermath of the Permo-Triassic mass

884 extinction: evidence from northern Italy. Palaeogeography, Palaeoclimatology, Palaeoecology

885 124: 137-–151.

886 Twitchett, R.J., and T. Oji. 2005. Early Triassic recovery of echinoderms. Comptes Rendus Palevol 4:

887 531-–542.

888 Urlichs, M. 1977. Zur Altersstellung der Pachycardientuffe und der Unteren Cassianer Schichten in

889 den Dolomiten (Italien). Mitteilungen der Bayerischen Staatssammlung für Paläontologie und

890 historische Geologie 17: 15-–25.

891 Urlichs, M. 1982. Zur Stratigraphie und Fossilführung des Lettenkeupers (Ob. Trias) bei Schwäbisch

892 Hall (Baden-Württemberg). Jahresberichte und Mitteilungen des Oberrheinischen Geologischen

893 Vereins, Neue Folge 64: 213-–224.

894 Urlichs, M. 1992. Vermeintlicher Zwergwuchs bei Muscheln aus dem untersten Trochitenkalk (Oberer

895 Muschelkalk, Mitteltrias) Württembergs. Carolinea 50: 9-–26.

896 Urlichs, M. 1994. Trachyceras LAUBE 1869 (Cephalopoda) aus dem Unterkarn (Obertrias) der Formatted: Font: Italic

897 Dolomiten (Italien). Stuttgarter Beiträge zur Naturkunde (B) 217: 1-–55.

898 Urlichs, M. 2014. A phylogenetic lineage of Myophoria kefersteini subspecies (Bivalvia) from the Late Formatted: Font: Italic

899 Ladinian to the Early Carnian (Triassic) in the Dolomites and Julian Alps (Italy). Neues Jahrbuch für

900 Geologie und Paläontologie, Abhandlungen 274: 187-–207.

901 Urlichs, M., and Tichy, G. 2000. Zur Alterstellung der Schlernplateau-Schichten (Mitteltrias) am

902 Schlern (westliche Dolomiten, Italien). Stuttgarter Beiträge zur Naturkunde (B) 297: 1-–10.

39

903 Velledits, F., C. Péró, J. Blau, B. Senowbari-Daryan, S. Kovács, O. Piros, T. Pocsai, H. Szügyi-Simon, P.

904 Dumitricã, and J. Palfy. 2011. The oldest Triassic platform margin reef from the Alpine-Carpathian

905 region (Aggtelek, NE Hungary): Platform evolution, reefal biota and biostratigraphic framework.

906 Rivista Italiana di Paleontologia e Stratigrafia 117: 221-–268.

907 Villier L., A. Brayard, K.G. Bylund, J.F. Jenks, G. Escarguel, N. Olivier, D.A. Stephen, E. Vennin, and E.

908 Fara. 2018. Superstesaster promissor gen. et sp. nov., a new starfish (Echinodermata, Asteroidea)

909 from the Early Triassic of Utah, USA, filling a major gap in the phylogeny of asteroids. Journal of

910 Systematic Palaeontology 16: 395-–415.

911 Volz, W. 1896. Die Korallen der Schichten von St. Cassian in Süd-Tirol (Frech, Fritz; Volz, Wilhelm: Die

912 Korallenfauna der Trias. Monographisch bearbeitet. II). Palaeontographica 43: 1-–124.

913 Vörös, A. 2003. The Pelsonian Substage on the Balaton Highland (Middle Triassic, Hungary).

914 Geologica Hungarica 55: 1-–52.

915 Waagen, L. 1903. Brachiopoden aus den Pachycardientuffen der Seiser Alpe. Jahrbuch der k. k.

916 geologischen Reichsanstalt 53: 443-–452.

917 Waagen, L. 1907. Die Lamellibranchiaten der Pachycardientuffe der Seiser Alm nebst vergleichend

918 paläontologischen und phylogenetischen Studien. Als Fortsetzung (II. Teil) zu A. Bittner,

919 Lamellibranchiaten der alpinen Trias. Abhandlungen der k. k. geologischen Reichsanstalt 18: 1-–

920 180.

921 Wagner, R. 1897. Beitrag zur genaueren Kenntnis des Muschelkalks bei Jena. Abhandlungen der

922 Königlich Preussischen Geologischen Landesanstalt Neue Folge 27: 1-–105.

923 Weidlich, O., W. Kiessling, and E. Flügel. 2003. Permian-Triassic boundary interval as a model for

924 forcing marine ecosystem collapse by long-term atmospheric oxygen drop. Geology 31: 961-–964.

40

925 Wignall, P.B., and R.J. Twitchett. 2002. Permian-Triassic sedimentology of Jameson Land, East

926 Greenland: incised submarine channels in an anoxic basin. Journal of the Geological Society

927 London 159: 691-–703.

928 Wilckens, R. 1909. Paläontologische Untersuchung triadischer Faunen aus der Umgebung von

929 Predazzo in Südtirol. Verhandlungen des Naturhistorisch-Medizinischen Vereins Heidelberg 10: 81-

930 –231.

931 Wöhrmann, S. von, and Koken, E. 1892. Die Fauna der Raibler Schichten vom Schlernplateau.

932 Zeitschrift der Deutschen geologischen Gesellschaft 44: 167-–223.

933 Wöhrmann, S. von. 1889. Die Fauna der sogenannten Cardita- und Raibler Schichten in den

934 Nordtiroler und Bayerischen Alpen. Jahrbuch der k. k. Geologischen Reichsanstalt 39: 181-–258.

935 Yang, H., Z.-Q. Chen, Y. Wang, W. Ou, W. Liao, and X. Mei. 2015. Palaeoecology of microconchids

936 from microbialites near the Permian-Triassic boundary in South China. Lethaia 48: 497-–508.

937 YingYin, H., and E.L. Yochelson. 1983a. Middle Triassic Gastropoda from Qingyan, Guizhou Province,

938 China: 1. Pleurotomariacea and Murchisoniacea. Journal of Paleontology 57: 162-–187.

939 YingYin, H., and E.L. Yochelson. 1983b. Middle Triassic Gastropoda from Qingyan, Ghizhou Province,

940 China: 3. Euomphalacea and Loxonematacea. Journal of Paleontology 57: 1098-–1127.

941 YingYin, H., and E.L. Yochelson. 1983c. Middle Triassic Gastropoda from Qingyan, Guizhou Province,

942 China: 2. Trochacea and Neritacea. Journal of Paleontology 57: 515-–538.

943 Yose, L.A. 1991. Sequence stratigraphy of mixed carbonate/volcaniclastic slope deposits flanking the

944 Sciliar (Schlern) – Catinaccio buildup, Dolomites, Italy. In The Northern Margin of the

945 Schlern/Sciliar-Rosengarten/Catinaccio Platform, eds. R. Brandner, E. Flügel, R. Koch, and L.A.

946 Yose, 17-–39. Dolomieu Conference on Carbonate Platforms and Dolomitization, Guidebook

947 Excursion A, Ortisei (Italy).

41

948 Zardini, R. 1978. Fossili Cassiani (Trias medio-superiore) – Atlante die gasteropodi della formazione di

949 S. Cassiano raccolti nella regiona dolomitica attorno a Cortina d’Ampezzo. Cortina d’Ampezzo:

950 Edizione Ghedina.

951 Zardini, R. 1981. Fossili Cassiani (Trias medio-superiore) – Atlante die bivalvi della formazione di S.

952 Cassiano raccolti nella regiona dolomitica attorno a Cortina d’Ampezzo. Cortina d’Ampezzo:

953 Edizione Ghedina.

954 Zatoń, M., H. Hagdorn, and T. Borszez. 2013a. Microconchids of the species Microconchus valvatus

955 (Münster in Goldfuss, 1831) from the Upper Muschelkalk (Middle Triassic) of Germany.

956 Palaeobiodiversity and Palaeoenvironments 94: 453-–461.

957 Zatoń, M., P.D. Taylor, and O. Vinn. 2013b. Early Triassic (Spathian) post-extinction Microconchids

958 from Western Pangea. Journal of Paleontology 87: 159-–165.

42

959 Figure captions

960

961 Fig. 1 Locality map showing the position of the studied sections at the Seiser Alm/Alpe di Siusi (red

962 stars) and the Schlern/Sciliar (blue stars). GPS-coordinates are N 46°30’40.8’’, E 11°33’48.54’’ for

963 Schlern locality A and N 46°30’36.19’’, E 11°33’51.9’’ for Schlern locality B

964

965 Fig. 2 Field photographs. (a) Sampling locality Frommerbach A, showing the investigated section (see

966 also Fig. 10 [left]). Scale = 1 m. (b) Sampling locality Frommerbach B, showing the investigated

967 section (see also Fig. 10 [right]). (c) Western flank of the Seiser Klamm/Gola di Siusi showing the

968 Schlernplateau beds (SchPl. beds), the post-volcanic bedded facies of the Rosszähne Fm. (RZ),

969 volcanic rocks (V) and the pre-volcanic Schlerndolomite of the Rosengarten Fm. (R). The white

970 rectangle indicates the position of (d). (d) Detailed look on the Schlernplateau beds. (e) Bioclastic

971 rudstone from top of the Schlernplateau beds containing gastropods and bivalves. (f) Oncolites from

972 the western Schlernplateau. Coin (16 mm) for scale

973

974 Fig. 3 Section of the Schlernplateau beds from the western flank of the Seiser Klamm/Gola di Siusi. M

975 = marl, m = mudstone, w = wackestone, p = packstone, g = grainstone, SP = Schlernplateau. The

976 Ladinian/Carnian boundary lies within bed SP10

977

978 Fig. 4 Rarefaction analyses from the Schlernplateau beds (a) and the Pachycardientuffe (b). Only

979 samples with more than 40 individuals were analysed. See text for details. SP = Schlernplateau, SK =

980 Seiser Klamm, SK-E = end of Seiser Klamm, FB = Frommmerbach

981

43

982 Fig. 5 Most common bivalves and gastropods collected at the Schlernplateau. (1) Trigonodus costatus

983 von Wöhrmann and Koken, 1892, left valve interior (1A) and left valve exterior (1B), PZO5984, bed

984 SP7. (2) Cassianella bipartita (von Klipstein, 1845), left valve, PZO5988, bed SP8. (3) Entolioides zitteli

985 (von Wöhrmann and Koken, 1892), right valve, PZO5986. (4) Myophoriopis richthofeni (Stur, 1868),

986 left valve, PZO5985, bed SP10. (5) Myophoria kefersteini (Münster in Goldfuss, 1837), left valve,

987 PZO5987, bed SP8. (6) ‘Loxonema’ lineatum von Wöhrmann and Koken, 1892, PZO5994, bed SP8. (7)

988 ‘Loxonema’ aequale von Wöhrmann and Koken, 1892, PZO5990, bed SP7. (8) Stephanocosmia

989 coronata (von Wöhrmann and Koken, 1892), PZO5989. (9) Zygopleura spinosa von Wöhrmann and

990 Koken, 1892, PZO5995bed SP8. (10) Pustularia alpina von Eichwald, 1851, PZO5998. (11) Tretospira

991 multistriata von Wöhrmann and Koken, 1892, PZO5997, bed SP8. (12) Coelostylina solida (von

992 Wöhrmann and Koken, 1892), PZO5996, bed SP7. (13) Neritaria plicatilis (von Klipstein, 1844),

993 PZO5993, bed SP8. (14) Naticopsis neritacea (Münster, 1841), PZO5991, bed SP7. (15) Palaeonarica

994 concentrica (Münster, 1841), PZO5992, bed SP7. Scale bars represent 1 cm for bivalves and 0.5 cm

995 for gastropods

996

997 Fig. 6 Bivalves from the Pachycardientuffe. (1) Pachycardia rugosa von Hauer, 1857, right valve

998 exterior (1A) and right valve interior (1B), PZO5980, bed FB B1. (2) Heminajas fissidentata (von

999 Wöhrmann, 1889), left valve, PZO5983, bed FB A3. (3) Pteria kokeni (von Wöhrmann and Koken,

1000 1892), left valve, PZO5982. (4) Pachycardia plieningeri Broili, 1903, left valve interior (4A) and left

1001 valve exterior (4B), PZO5981, bed FB B1. (5) Myoconcha parvula von Wöhrmann and Koken, 1892,

1002 right valve, SMNHS75537. (6) Pteria salomoni (Broili, 1903), right valve, SMNHS75534/1-2. (7)

1003 Myoconcha curvata Broili, 1903, left valve interior and left valve exterior, SMNHS10274. (8)

1004 Mysidioptera interrupta Broili, 1903, left valve, SMNHS17501/162. (9) Trigonodus rablensis (Gredler,

1005 1862), right valve exterior (9A) and left valve interior (9B), 10284/1-3. (10) Waagenoperna planata

1006 (Broili, 1903), left valve, SMNHS17501/45. (11) Mysidioptera emiliae Bittner, 1900, right valve,

1007 SMNHS17501/12. (12) Pteria tofanae (Bittner, 1895), left valve, SMNHS75538, collected at 44

1008 Tschapitbach. (13) Newaagia lata (von Klipstein, 1845), right valve interior, SMNHS75536. (14)

1009 Parallelodon tschapitana (Broili, 1903), right valve, SMNHS17501/250. Scale bars represent 1 cm

1010

1011 Fig. 7 Q-mode (samples) and R-mode (species) cluster analysis of samples with more than 40

1012 individuals from the Schlernplateau beds using unweighted pair-group average and Morisita

1013 similarity index. The circle sizes represent the abundance of species within the corresponding sample

1014 (small circle = 1 individual, small medium-sized circle = 2-5 individuals, big medium-sized circle = 6-20

1015 individuals, big circle = more than 20 individuals). SP = Schlernplateau, SK = Seiser Klamm, SK-E = end

1016 of Seiser Klamm. See text for details

1017

1018 Fig. 8 Rank-abundance distributions and guild-structures of the entire fauna of the Schlernplateau

1019 beds (a) and Pachycardientuffe (b). n = total number of individuals

1020

1021 Fig. 9 Rank-abundance distributions and guild-structures of representative fossiliferous beds SP14

1022 (a), SP7 (b) and SP3 (c) from the Schlernplateau beds (see also Fig. 3). n = total number of individuals

1023

1024 Fig. 10 Sections Frommerbach A and B of the Pachycardientuffe from the Seiser Alm/Alpe di Siusi

1025

1026 Fig. 11 Rank-abundance distributions and guild-structures from the fossiliferous beds FB A3 (a), FB B2

1027 (b) and FB B1 (c) from the Pachycardientuffe (see also Fig. 10). n = total number of individuals

1028

45

1029 Fig. 12 Lithological samples from locality Frommerbach A showing the highly fossiliferous limestone

1030 (a) and volcanoclastic sandy siltstone (b-c) containing disarticulated, randomly arranged bivalves.

1031 Scales are 23 mm (diameter of the coin) for (a) and 5 cm for (b) and (c)

1032

1033 Fig. 13 (a) Palaeoenvironmental reconstruction of the Schlern and Seiser Alm area during the Late

1034 Ladinian showing the distribution of sedimentary facies and the potential source areas of the

1035 Pachycardientuffe. (b) Panorama image taken from the top of the eastern margin of the Schlern

1036 mountain, showing present-day position of the former lagoon (Schlernplateau beds, 1), the post-

1037 volcanic reef (Rosszähne Fm., 2), the probable position of the suggested volcanic island (3) and the

1038 basin (Seiser Alm basin, Pachycardientuffe and Wengen Fm., 4)

1039

1040 Fig. 14 Diversity changes throughout the Early and Middle Triassic based on data shown in Tab. 1; see

1041 text for further explanation. (a) Total number of species. (b) Gastropod/bivalve-ratio. Blue lines mark

1042 formations with carbonate platform facies. G = Griesbachian; D = Dienerian; Sm = Smithian; Sp =

1043 Spathian, A = Aegean; B = Bithynian; P = Pelsonian; I = Illyrian; F = Fassanian; L = Longobardian; J =

1044 Julian; Carn. = Carnian; LT = Late Triassic. Marmolada = Marmolada Limestone; PachyT =

1045 Pachycardientuffe; Latemar = Latemar Limestone; SchPl = Schlernplateau beds

46

1046 Table captions

1047

1048 Tab. 1 Benthic species richness of selected Early and Middle Triassic faunas. B = bivalves, G =

1049 gastropods, Br = brachiopods, C = corals, Sp = sponges, Cr = crinoids, E = echinoids, A = asteroids, Bry

1050 = bryozoans, Cru = crustaceans, H = holothurioids, M = microconchids, O = ophioroids, Pc =

1051 polychaete worms, Pp = polyplacophorans, Sc = scaphopods, ? = these taxa were reported but the

1052 exact number of species is unknown. Information about the age of the lithological units come from

1053 Fürsich and Wendt 1977, Broglio Loriga et al. 1990, Brack and Rieber 1993, Brack et al. 1996, Vörös

1054 2003, Manfrin et al. 2005, Stiller and Bucher 2008, Hagdorn and Nitsch 2009, Shigeta et al. 2009,

1055 Balini et al. 2010, Hofmann et al. 2013a, 2013b, 2014, 2015, Urlichs 2014, Foster et al. 2015, 2017,

1056 Hausmann and Nützel 2015, Hautmann et al. 2015, Brayard et al. 2017, Foster and Sebe 2017. See

1057 text for details

47

1058 Online Resources captions

1059

1060 Online Resource 1 Excel tables showing the quantitative matrix of fossil occurrences used for data

1061 analyses, the comparison between the faunas of the Schlernplateau beds and the Pachycardientuffe

1062 as well as the number of species, number of individuals and Simpson-D values for each fossiliferous

1063 bed

48

Figure Click here to download Figure Fig1.tif Figure Click here to download Figure Fig2.tif Figure Click here to download Figure Fig3.tif Figure Click here to download Figure Fig4.tif Figure Click here to download Figure Fig5.tif Figure Click here to download Figure Fig6.tif Figure Click here to download Figure Fig7.tif Figure Click here to download Figure Fig8.tif Figure Click here to download Figure Fig9.tif Figure Click here to download Figure Fig10.tif Figure Click here to download Figure Fig11.tif Figure Click here to download Figure Fig12.tif Figure Click here to download Figure Fig13.tif Figure Click here to download Figure Fig14.tif Table

number of species total number others (A, lithological unit age of benthic Bry, Cru, H, reference(s) B G Br C Sp Cr E species M, O, Pc, Pp, Sc) Laube (1865a-b, 1868, 1869a), Kittl (1891, 1892, 1894), Bittner 5 H (1890, 1892, 1895), Volz (1896), 4 Sc Pia (1937), Leonardi and Fiscon Cassian Fm. Longobardian – 1146 296 525 110 72 79 15 34 3 O (1947, 1959), Frizzell and Exline (Italy) Julian 2 Bry (1955), Fürsich and Wendt 1 Pc (1977), Zardini (1978, 1981), Nützel and Kaim (2014), Hausmann and Nützel (2015) von Wöhrmann and Koken Schlernplateau beds Late Longobardian – 85 27 47 5 ? 2 4 (1892), Blaschke (1905), this (Italy) Early Julian study Broili (1903), Blaschke (1905), Pachycardientuffe Late Longobardian 277 150 77 24 5 3 4 13 1 Sc Waagen (1903, 1907), Fürsich (Wengen Fm., Italy) and Wendt (1977), this study Erfurt Fm. 2 Cru Schmidt (1928, 1938), Urlichs Early Longobardian 78 47 23 5 (“Lower Keuper”, Germany) 1 Pc (1982) 14 Cru Wagner (1897), Schmidt (1928, 9 O Meissner Fm. 1938), Busse (1972), Hagdorn Late Illyrian Early 5 Pc – 250 118 79 10 3 1 2 5 and Mundlos (1982), Hagdorn (upper part of the „Upper Longobardian 2 Sc Muschelkalk and Ockert (1993), “, Germany) 1 A Zatoń et al. (2013a) 1 M Middle Illyrian Marmolada Limestone – Middle 290 45 201 31 7 2 3 1 Cru Böhm (1895), Salomon (1895) (Italy) Longobardian Latemar Limestone Middle Illyrian Early – 125 24 86 15 Häberle (1908), Wilckens (1909) (Italy) Longobardian 10 Cru Wagner (1897), Schmidt (1928, 9 O 1938), Busse (1972), Hagdorn Trochitenkalk Fm. 5 Pc and Ockert (1993), Ockert (basal part of the Middle Illyrian 209 96 70 5 2 1 2 5 “Upper 2 Sc (1993), Urlichs (1992),

Muschelkalk”, Germany) 1 A Hautmann and Hagdorn (2013), 1 M Zatoń et al. (2013a) Karlstadt Fm., Heilbronn Fm., Diemel Fm. 2 Pc Early Illyrian 81 34 38 3 1 ? 1 1 Schmidt (1928, 1938) („Middle Muschelkalk“, 1 Cru Germany) 4 Bry Yin and Yochelson (1983a-c), Leidapo Mb. Late Pelsonian – 308 109 126 27 8 3 8 17 3 Sc Stiller (2001), Komatsu et al. (Quingyan Fm., China) Early Illyrian 3 V (2004) 1 M Zuhánya Limestone Late Pelsonian – 33 19 5 2 3 1 1 O Foster and Sebe (2017) (Hungary) Early Illyrian 1 Sc Szabó et al. (1979), Vörös Felsőörs Fm. Pelsonian Illyrian 77 39 35 1 2 H (Hungary) – (2003) 8 Cru 5 Pc Jena Fm. Late Bithynian Early 4 O Wagner (1897), Schmidt (1928, – 263 115 93 11 7 1 12 1 (“Lower Muschelkalk“, Illyrian 4 Sc 1938) Germany) 1 A 1 Pp 1 M Lapis Limestone Late Bithynian Early – 23 12 3 2 3 1 O Foster and Sebe (2017) (Hungary) Pelsonian 1 Sc Röt Fm. Wagner (1897), Schmidt (1928, Early Bithynian 59 38 3 2 1 1 14 Cru (“Upper Buntsandstein“, 1938) Germany) Viganvár Limestone Late Aegean Early 1 M – 20 10 5 2 1 Foster and Sebe (2017) (Hungary) Bithynian 1 Sc Csopak Marl Spathian 19 15 3 1 ? ? O Broglio Loriga et al. (1990), (Hungary) Budai (2008) McGowan et al. (2009), Virgin Fm. Hautmann et al. (2013), Spathian 46 30 6 5 1 2 1 1 M (USA) Hofmann et al. (2013b), Zatoń et al. (2013b) Szinpetri Limestone Fm. 1 M Late Spathian 11 6 2 1 Foster et al. (2015) (USA) 1 O Neri and Posenato (1985), San Lucano Mb. Late Spathian 12 7 4 1 Broglio Loriga et al. (1990), (Werfen Fm., Italy) Foster et al. (2017) Neri and Posenato (1985), ? M Cencenighe Mb. Broglio Loriga et al. (1990), Middle Spathian 34 25 6 1 1 O (Werfen Fm, Italy) Hofmann et al. (2015), Foster et 1 Sc al. (2017) 1 M Szin Marl Fm. Early Middle – 18 10 4 1 1 ? O Foster et al. (2015) (Hungary) Spathian 1 Sc Broglio Loriga et al. (1990), Val Badia Mb. ? M Early Spathian 23 15 5 1 1 Hofmann et al. (2015), Foster et (Werfen Fm., Italy) 1 O al. (2017) Lower Shale Unit Late Smithian Early 2 Cru – 14 5 3 2 1 Brayard et al. (2017) (Thaynes Gr., USA) Spathian 1 O 1 A Batten and Stokes (1986), Sinbad Fm. ? Sc Nützel and Schulbert (2005), Smithian 48 18 27 2 ? ? (USA) ? O Brayard et al. (2011), Hofmann ? Pc et al. (2014), Villier et al. (2018) Hidegkút Fm. Bittner (1901), Broglio Loriga et Smithian 16 15 1 (Hungary) al. (1990) Broglio Loriga et al. (1990), Campil Mb. Smithian 26 21 3 1 Hofmann et al. (2015), Foster et (Werfen Fm., Italy) al. (2017) Zhitkov Fm. ? M Dienerian Smithian 22 11 8 1 1 Shigeta et al. (2009) (Russia) – ? O 1 Sc Broglio Loriga et al. (1990), Gastropod Oolite Late Dienerian – 16 11 4 1 ? M Hofmann et al. (2015), Foster et (Werfen Fm., Italy) Early Smithian al. (2017) ? M Broglio Loriga et al. (1990), Seis Mb. Dienerian 24 18 4 1 ? ? O Hofmann et al. (2015), Foster et (Werfen Fm., Italy) 1 Sc al. (2017) Dinwoody Fm. Griesbachian – 19 15 2 2 Hofmann et al. (2013a) (USA) Dienerian Köveskál Fm., Arács Fm., Griesbachian Alcsútdoboz Fm. – 20 12 7 1 Broglio Loriga et al. (1990) Dienerian (Balaton Highland, Hungary) Lazurnaya Bay Fm. Griesbachian 26 12 6 8 Shigeta et al. (2009) (Russia) Middle Bódvaszilas Sandstone Fm. Griesbachian 14 12 1 1 M Foster et al. (2015) (Hungary) – Smithian Luolou Fm. Middle ? M 30 20 5 3 ? ? Hautmann et al. (2015) (China) Griesbachian ? Pc Andraz Horizon Middle 3 3 Foster et al. (2017) (Werfen Fm., Italy) Griesbachian Broglio Loriga et al. (1986, Mazzin Mb. Early Middle ? M – 17 12 3 1 1990), Hofmann et al. (2015), (Werfen Fm., Italy) Griesbachian 1 Pc Foster et al. (2017) Late Tesero Oolite ? M Broglio Loriga et al. (1986, Changhsingian 15 5 3 5 ? 1 (Werfen Fm., Italy) – 1 Pc 1990), Foster et al. (2017) Early Griesbachian

Supplementary Material

Click here to access/download Supplementary Material ESM_1.xlsx