1 Crustaceans in cold seep ecosystems: fossil record, geographic distribution, taxonomic composition, 2 and biology 3 4 Adiël A. Klompmaker1, Torrey Nyborg2, Jamie Brezina3 & Yusuke Ando4 5 6 1Department of Integrative Biology & Museum of Paleontology, University of California, Berkeley, 1005 7 Valley Life Sciences Building #3140, Berkeley, CA 94720, USA. Email:
[email protected] 8 9 2Department of Earth and Biological Sciences, Loma Linda University, Loma Linda, CA 92354, USA. 10 Email:
[email protected] 11 12 3South Dakota School of Mines and Technology, Rapid City, SD 57701, USA. Email: 13
[email protected] 14 15 4Mizunami Fossil Museum, 1-47, Yamanouchi, Akeyo-cho, Mizunami, Gifu, 509-6132, Japan. 16 Email:
[email protected] 17 18 This preprint has been submitted for publication in the Topics in Geobiology volume “Ancient Methane 19 Seeps and Cognate Communities”. Specimen figures are excluded in this preprint because permissions 20 were only received for the peer-reviewed publication. 21 22 Introduction 23 24 Crustaceans are abundant inhabitants of today’s cold seep environments (Chevaldonné and Olu 1996; 25 Martin and Haney 2005; Karanovic and Brandão 2015), and could play an important role in structuring 26 seep ecosystems. Cold seeps fluids provide an additional source of energy for various sulfide- and 27 hydrocarbon-harvesting bacteria, often in symbiosis with invertebrates, attracting a variety of other 28 organisms including crustaceans (e.g., Levin 2005; Vanreusel et al. 2009; Vrijenhoek 2013). The 29 percentage of crustaceans of all macrofaunal specimens is highly variable locally in modern seeps, from 30 0–>50% (Dando et al. 1991; Levin et al.