Research articleUnravelling the paleoecology of flat clams: New insights from an Upper Triassic halobiid bivalve
Introduction
Flat clams are bivalves that are characterized by extremely thin shells and low valves convexity. They were ubiquitous in rocks of Devonian to Jurassic age (Jefferies and Minton, 1965). Sporadically found in (possibly) shallower environments (Miroshnikov and Burii, 1969; Hatleberg and Clark, 1984; Ando, 1987; Campbell, 1994), they thrived in deep-water deposits where they often form the main biogenic component. During the Middle-Upper Triassic, genera such as Daonella and Halobia were incredibly widespread. This fact, along with a fast species turnover, has made them powerful biostratigraphic tools, with an average species duration similar to that of coeval ammonoid species (e.g. around 1–2 Ma: McRoberts, 2010).
Since the early works of von Mojsisovics, 1869, von Mojsisovics, 1874, flat clams-bearing sedimentary beds of different ages have drawn the attention of a great number of researchers with different backgrounds. Paleontologists, geochemists and sedimentologists have investigated the peculiar distribution of these organisms. However, environmental interpretations often diverge dramatically. Integrated and multi-approach studies investigating the paleoecology of these organisms are lacking, and, too often, interpretations have been made taking into account only a few sets of parameters.
Through a multi-disciplinary approach, this work aims to better characterize the paleoecology, lifestyle and habitat of these conspicuously flat, deep-water Triassic-Jurassic bivalves, highlighting their potential for paleo-environmental reconstruction. We initially provide a summary of current understanding of the mode of life of “flat clams” (Fig. 1) from different ages, with a focus on mostly Triassic and Jurassic forms, which are summarized, discussed, and compared to possible extant counterparts. The second part of the manuscript is a case study of an Upper Triassic, Halobia-rich, deep-water succession exposed near Tsiko Lake, Vancouver Island, Canada. There, the sequence was investigated by means of a multi-disciplinary approach including sedimentology, taphonomy, morphometry, microscopy, and geochemistry.
Section snippets
Living on, near, or in the seabed: Benthic
Several authors see the lowered volume/surface ratio of flat clams as an adaptation to soft substratum and low oxygen levels in bottom waters (Rhoads and Morse, 1971; Savrda et al., 1984; Thompson et al., 1985; Fürsich et al., 1991; Kelly and Doyle, 1991; Savrda and Bottjer, 1991; Hollingworth and Wignall, 1992; Wignall, 1993; Wignall and Pickering, 1993; Wignall, 1994; Etter, 1995, Etter, 1996; Röhl et al., 2001; Harzhauser and Mandic, 2004; Waller and Stanley, 2005; Schatz, 2005; Bakke, 2017;
Geological context
The bedrock geology of Vancouver Island is principally formed by a thick Late Paleozoic to Early Mesozoic volcano-sedimentary succession regarded as being part of the tectonostratigraphic Wrangellia terrane (Jones et al., 1977). Wrangellia, which stretches north into southern Alaska via the Haida Gwaii Islands (Wheeler and McFeeely, 1991), is inferred to have collided with the Alexander terrane to form the Insular belt in the Late Carboniferous (Gardner et al., 1988), and merged to inboard
Depositional environments
The succession comprises sediments deposited in a slope to basinal setting, above the CCD (or ACD if the inner layer of Halobia was originally made of aragonite as suggested by Carter, 1990), in a low-energy environment, well below the storm wave base, with disturbance only at irregular times by the gravitational re-deposition of sediments reworked from the outer platform. These alternations of hemipelagic sediments and platform to slope-derived sediments are similar to the depositional
Conclusions
Flat clams-bearing hemipelagic litho-biofacies at Tsiko Lake were deposited under generally restricted oxygen conditions. The entire halobiid assemblage is monospecific and consists of specimens of Halobia cordillerana: the abundance and dimensions (represented by their shell thickness) of these halobiids correlate with bottom water redox conditions inferred from redox proxies and paleontological and ichnofacies data. Based on different lines of evidence, we suggest an epibenthic recliner mode
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.
Acknowledgments
Special thanks to Christopher McRoberts for his valuable help with the fossil identifications. We thank the lab staff at University of Lausanne and in particular Thierry Adatte for the help with Rock-Eval analyses. We thank Manuel Rigo and another anonymous reviewer for their useful comment which improved this manuscript. The authors acknowledge Stephen Reid, Andrew Hobson and Yijun Xiong for the help with geochemical analyses, and François Gischig for the great quality of the thin sections
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