Amano et al.—New and Mesozoic-relict mollusks from Paleocene wood-fall communities
foraminifers show a lower extinction rate than shallow-water ones in the end-Cretaceous extinction because of little environ- mental change (Kaiho, 1994). Consequently, the low extinction rate of protobranchs is attributed to their deposit feeding strategy and preference of dwelling in deep water (e.g., Dame, 1996). Five of the oldest fossil records of Recent molluscan genera
have been found in the Katsuhira Formation: Tindaria Bellardi, 1875, Bentharca, Poromya, Neverita,and Admete Krøyer in Möller, 1842 (Amano et al., 2015a, 2016b; Amano and Jenkins, 2017; this study; Table 8). Moreover, Urahorosphaera kanekoi Amano and Oleinik, 2014 is the one of the oldest records of buccinoids in the Paleocene (Amano and Oleinik, 2014). Some Recent genera of deep-sea chemosynthetic bivalves also origi- nated in the North Pacificafter the middle Eocene: Vulcanidas Cosel and Marshall, 2010, Bathymodiolus Kenk and Wilson, 1985, Calyptogena Dall, 1891, and Archivesica Dall, 1908 (Kiel and Amano, 2010, 2013; Amano and Kiel, 2010; Amano et al., 2014). It is interesting to note that species of these genera are epifaunal or very shallowly infaunal, regardless of feeding type. However, the exact reason why these Recent genera originated in the North Pacific is unknown. From these, although the Katsuhira fauna includes many Mesozoic-relict taxa as refu- gees, some Cenozoic species first appeared in this formation. From the viewpoint of the evolution of wood-fall commu-
nities, the fauna of the Katsuhira Formation includes three chemosynthesis-based species: the limpet Bathyacmaea? sp., the lucinid Myrtea ezoensis, and the thyasirid Thyasira oliveri n. sp. Consequently, the Paleocene wood-fall communities from the Katsuhira Formation in Hokkaido and the Basilika Forma- tion in Spitsbergen have no provannid gastropods and no small mussels, compared to the first appearance of provannids in the Upper Cretaceous wood-fall communities (Kiel et al., 2009). Based on the protobranch species from the Katsuhira Formation (Amano and Jenkins, 2017), the paleobathymetry of the for- mation can be estimated to be between 200 and 500m in depth. According to Hryniewicz et al. (2016), the Basilika Formation was deposited in an offshore prodelta. On the other hand, most Recent species of Provanna Dall, 1918 live deeper than 500m (see Amano and Little, 2014, table 1). Because the Paleocene wood-fall communities lived in shallower water than the habitat of provannids, they lack provannid gastropods. On the other hand, despite the fact that small mussels can live in shallower water (e.g., Lorion et al., 2010), they have not been found in the Paleocene sites and even in the early to middle Eocene wood- fall communities from Washington State in the US (Kiel, 2008). The oldest record of small mussels from the wood-fall com- munities has been found from the upper Eocene part of the Lincoln Creek Formation (Kiel and Goedert, 2006). Adding to the previous Paleocene fossil records, this study also confirms that small bathymodioline mussels in the chemosynthetic communities did not appear in the Paleocene.
Acknowledgments
We thank A.G. Beu (GNS Science) for his critical reading of the manuscript and useful suggestions. We also thank K. Inoue (Obihiro City) for kindly offering his collection, and H. Nishi and J. Nemoto (Tohoku University), Y. Kobayashi (Hokkaido University), and T. Haga (National Museum of Nature and
645
Science, Tokyo) for their help to examine the type material.We also thank A. Kaim (Institute of Paleobiology, Polish Academy of Sciences) and A. Oleinik (Florida Atlantic University) for their reviewof this paper and many useful comments. This study was partly supported by a Grant-in-aid for Scientific Research from the Japan Society for the Promotion of Science (C, 26400500, 2014−2016; C, 17K05691, 2017 −2019) to KA and RGJ, and a Grant for Program to Disseminate Tenure Tracking System (JST) to RGJ.
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