Major coral extinctions during the early Toarcian global warming event
Introduction
The history of life is punctuated by extinction events that drove evolutionary dynamics, and scientists posit that the Earth is currently entering a similar mass extinction (Ceballos et al., 2015). Five major “first-order” biotic crises are reported for the Phanerozoic, followed by “second-order” crises, such as the Pliensbachian-Toarcian crisis at ca. 183 Ma (e.g., Raup and Sepkoski, 1982; Bambach et al., 2004). The distinction between first and second-order crises remains unclear as the ranking of crises varies according to the considered characters, such as taxonomic scale (i.e., families, genera, species), functional niche loss, consideration of associated palaeoenvironmental upheaval, and the different ways to statistically weigh raw data (Benton, 1995).
The Pliensbachian-Toarcian (Pl-To) transition was marked by drastic and repeated palaeoenvironmental disturbances associated with the emplacement of the Karoo-Ferrar igneous province in southern Gondwana (Burgess et al., 2015). These environmental changes included major shifts in the temperature, chemistry, and oxygenation of the oceans, culminating in the Toarcian Oceanic Anoxic Event (T-OAE) (e.g.,Suan et al., 2010; Gómez and Goy, 2011; Caruthers et al., 2011; Dera et al., 2011; Reolid et al., 2020). Following the middle Pliensbachian warm period (Gómez et al., 2016), the latest Pliensbachian is marked by a significant cooling event with important sea-level fluctuations (De Graciansky et al., 1998; Aurell et al., 2003; Suan et al., 2010; Pittet et al., 2014; Krencker et al., 2019). The Pl-To boundary coincides with a pronounced warming event (Suan et al., 2010), enhanced continental weathering, and a widespread demise of neritic carbonate ecosystem (Bodin et al., 2016; Krencker et al., 2020), which often led to strong condensation of the sedimentary record or even a hiatus in sedimentation (Morard et al., 2003; Léonide et al., 2012; Krencker et al., 2014; Pittet et al., 2014). Climate cooling is observed during the remainder of the earliest Toarcian (Polymorphum zone; Ruebsam et al., 2019; Krencker et al., 2020), allowing renewed neritic carbonate production in several areas across the Tethys Ocean (Léonide et al., 2012; Trecalli et al., 2012; Krencker et al., 2020). At the onset of the T-OAE, a second demise of neritic carbonate ecosystems occurred, likely triggered by a profound carbon cycle perturbation, soaring global temperature (Suan et al., 2010; Korte et al., 2015), and a second pulse of continental weathering (Bodin et al., 2010; Krencker et al., 2020; Ettinger et al., 2021). Neritic carbonate production resumed along the southern Tethyan margin during the recovery of the carbon isotope excursion associated with the T-OAE (Trecalli et al., 2012; Krencker et al., 2014, Krencker et al., 2015) but not until at least the middle Toarcian along the northern Tethyan margin (Blomeier and Reijmer, 1999; Aurell et al., 2003; Léonide et al., 2012; Brigaud et al., 2014).
The Pl-To transition is one of the five most significant reefal crises in the geological record (Flügel and Kiessling, 2002; Kiessling and Simpson, 2011; Kiessling, 2011) and this event is correlated with significant extinctions across a wide number of marine taxa (Little and Benton, 1995; Ruban, 2004; Mattioli et al., 2009; Dera et al., 2010; Caruthers et al., 2013; Brame et al., 2019) as well as continental plants (Slater et al., 2019). The existing literature on corals during the Pl-To transition was highly dependent on the taxonomic work of Beauvais (1986). Indeed, previous studies (Kiessling et al., 2007; Lathuilière and Marchal, 2009) identified the need for more original field data followed by a revision of the taxonomy based on these new specimens and with consideration of possible intraspecific variability. This study provides a stronger analysis of Pliensbachian and Toarcian coral assemblages based in extensive new collections from Morocco (Middle and High Atlas), Italy (Calabria), France (Calvados and Pyrenees) and a deep revision of taxa from the literature (Vasseur, 2019; Vasseur and Lathuilière, 2020 in press). It also offers the first opportunity to examine coral assemblages in between the Pliensbachian/Toarcian stage boundary and T-OAE.
Section snippets
Material
Data have been gathered by the authors from five field missions in Morocco (Dadès Valley, Amellagou, Ziz Valley and Guigou Plateau) and one in Italy (Caloveto, Calabria). In addition to corals, ammonites were also collected to improve the age calibrations for this material using the most recent revisions of ammonite determination and biozones (Dommergues and Meister, 2017). New specimens collected by Marc Chesnier from Normandy (France) and Philippe Fauré from the French Pyrenees were also
Pliensbachian coral assemblages
In the field, Pliensbachian coral faunas are widely dominated in both diversity and quantity by solitary and phaceloid forms, while highly integrated forms (i.e., plocoid, cerioid, thamnasterioid, and meandroid) are rare and often occur as diminutive colonies (Table 1, Fig. 1). One exception to this general trend is in the Middle-Atlas and Calabria, where large, high-integration corals occur with higher diversity and skeletal volume in quite distal position.
In proximal environments, corals
Discussion
The results presented here illustrate the importance of the Pl-To crisis in coral communities; the Pliensbachian – Toarcian crisis marks the disruption between an early period of high disparity (represented by the amount of Pliensbachian families) and the later diversification of surviving taxa during the Toarcian and later. Furthermore, the Pliensbachian and post-T-OAE intervals mark the development of coral pioneers in the colonisation of deeper environments, with many specimens recovered
Conclusion
This study highlights the significance of both the Pl/To boundary and the T-OAE crises in coral evolution. These events resulted in extinction rates as high as 51.1% at the generic level and 90.9% at the species level, and combined (i.e., the Pl-To interval) rank as one of the three most important extinction and recovery periods in the geological history of Scleractinia, on par with (or even more devastating than) the T/J and K/Pg extinctions. Although it is not considered a first order biotic
Declaration of Competing Interest
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
The authors declare the following financial interests/personal relationships which may be considered as potential competing interests.
Acknowledgements
We are indebted to a number of people who took part in the field work, collection of specimens and work in museums. For our five field missions in Morocco we thank Hamid Haddoumi, Khalid Elhmidi, Mohamed Ousri, Rossana Martini, Simon Boivin, Hassan El Arabi, André Charrière, Thomas Saucède, Pierre Pellenard, Nicholas Ettinger, and Hannah Brame. We are especially thankful to Philippe Fauré who collected corals from the Pyrenees and who waited so long for the results. We thank Massimo Santantonio
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