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Miocene Abyssochrysoid Gastropod Provanna from Japanese Seep and Whale-Fall Sites

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Miocene Abyssochrysoid Gastropod Provanna from Japanese Seep and Whale-Fall Sites Miocene abyssochrysoid gastropod Provanna from Japanese seep and whale-fall sites KAZUTAKA AMANO and CRISPIN T.S. LITTLE Amano, K. and Little, C.T.S. 2014. Miocene abyssochrysoid gastropod Provanna from Japanese seep and whale-fall sites. Acta Palaeontologica Polonica 59 (1): 163–172. We describe three Miocene species of Provanna from Japan, two new and one in open nomenclature, that represent the only known fossil examples from whale-falls and a considerable increase in the Miocene diversity of the genus. Provanna hirokoae sp. nov. comes from the latest Middle Miocene Kuroiwa seep site in central Honshu. The shells of this species are mostly recrystallized, but contain relict crossed lamellar microstructures. Provanna alexi sp. nov. is from the early Middle Miocene Shosanbetsu whale-fall site in northwestern Hokkaido, and has well preserved shells comprising an outer simple prismatic layer and an inner crossed lamellar layer. The two Provanna specimens from the Middle Miocene Rekifune whale-fall site, in eastern Hokkaido, are preserved as external moulds only, so are left in open nomenclature. Based on current knowledge, the presence of an outer prismatic layer and an underlying crossed lamellar layer seems to be a common feature in the shells of Provanna, as well as in other genera belonging to the family Provan- nidae and the superfamily Abyssochrysoidea. Although the oldest occurrence of Provanna was in the Late Cretaceous, the genus did not spread geographically and ecologically until the Miocene (with four, or possibly five species), a date concordant with some molecular estimates. However, this could be an artefact of the fossil record because the known pre-Miocene seep and whale-falls are more geographically restricted than those from the Miocene. Key words: Mollusca, Gastropoda, Provanna, seep, whale-fall, Miocene, Japan. Kazutaka Amano [[email protected]], Department of Geoscience, Joetsu University of Education, 1Yamayashiki, Joetsu 943-8512, Japan; Crispin T.S. Little [[email protected]], School of Earth and Environment, University of Leeds, Leeds LS2 9JT, UK. Received 6 January 2012, accepted 24 May 2012, available online 5 June 2012. Copyright © 2014 K. Amano and C.T.S. Little. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Japan, one from upper Eocene to Oligocene seep deposits Introduction and upper Eocene wood-falls in Washington State, USA, and one from Miocene seeps in New Zealand (Table 1). In addi- The small gastropod genus Provanna Dall, 1918 is one of the tion to these, fossil gastropods from four different Miocene characteristic and species-rich molluscan taxon in modern localities have been figured, but not formally described, as chemosynthetic faunas (e.g., Warén and Bouchet 1993, 2001, Provanna or other provannids. These include “provannids” 2009; Sasaki et al. 2010). Most Provanna species are graz- from the Freeman’s Bay Limestone (a suspected seep depos- ers on filamentous bacteria, while some are detritus feeders. it) of the Miocene Lengua Formation, Trinidad (Gill et al. Although Bergquist et al. (2007) suggested that Provanna 2005: fig. 7I, J) and three Provanna species from two whale- variabilis Warén and Bouchet, 1986 may also harbor sym- fall sites and one seep site from Japan (Amano and Little biotic bacteria, Sasaki et al. (2010) raised doubts about this 2005; Amano et al. 2007, 2010). Here we formally describe interpretation based on the anatomy of the species. these Japanese species and discuss the shell microstructures Eighteen Recent species of Provanna have so far been and the fossil record of the genus Provanna. The Japanese described (Table 1), most from hydrothermal vent and hy- occurrences represent the only known fossil examples of drocarbon (cold) seep sites, but some also from whale-fall Provanna from whale-falls and a significant increase in the and wood-fall sites (e.g., Smith and Baco 2003; Warén and Miocene species diversity of the genus. Bouchet 2001, 2009; Sasaki et al. 2010). In contrast to this di- versity, only four fossil Provanna species have been formally Institutional abbreviations.—JUE, Joetsu University of Edu- described: two from Cretaceous seep deposits in Hokkaido, cation, Joetsu, Japan. Acta Palaeontol. Pol. 59 (1): 163–172, 2014 http://dx.doi.org/10.4202/app.2012.0002 164 ACTA PALAEONTOLOGICA POLONICA 59 (1), 2014 130° 140° one specimen of P. reticulata Warén and Bouchet, 2009. This was collected with samples of seep carbonate (M56B, GeoB 8212-1 TV grab) from 3100 m water depth at the Hydrate 1 Hole pockmark site, on the Congo deep-sea fan (Sahling et al. 2008) during RV METEOR Cruise M56 (Chief Scientist 2 Gerhard Bohrmann, MARUM, University of Bremen, Ger- many). 40° Systematic paleontology Phyllum Mollusca Linnaeus, 1758 Japan Sea 3 Class Gastropoda Cuvier, 1797 Pacific Ocean Order Caenogastropoda Cox, 1959 Superfamily Abyssochrysoidea Tomlin, 1927 Family Provannidae Warén and Ponder, 1991 Genus Provanna Dall, 1918 Type species: Trichotropis (Provanna) lomana Dall, 1918; Recent, US 30° Pacific Coast. Provanna hirokoae sp. nov. Fig. 2. 2010 Provanna sp.; Amano et al. 2010: figs. 5C–E. Etymology: Named for the first author’s wife Hiroko. Fig. 1. Localities of Miocene Provanna in Japan. 1, Shosanbetsu whale-fall Type material: Holotype, JUE 15901 (Fig. 2A); Paratypes, JUE 15902- site; 2, Rekifune whale-fall site; 3, Kuroiwa hydrocarbon seep site. 1 (Fig. 2B), JUE 15902-2 (Fig. 2C), JUE 15902-3 (Fig. 2D), JUE 15902-4 (Fig. 2E). Type locality: Kita-Kuroiwa Quarry, Kakizaki-ku, Joetsu City in central Material Honshu, Japan (see Amano et al. 2010); seep site. Type horizon: Fossil hydrocarbon seep deposits of the Ogaya Forma- Twenty-seven Provanna specimens were collected from the tion, uppermost Middle Miocene. Shosanbetsu whale-fall site in northwestern Hokkaido (local- Dimensions.—See Table 2. ity 1 in Fig. 1; Amano and Little 2005). This site is from the lower Middle Miocene (15.9–14.9 Ma) Chikubetsu Forma- Diagnosis.—Medium-sized Provanna with variable sculp- tion (Amano et al. 2007). We examined and measured nine ture ranging from sigmoidal axial growth lines and many of the specimens, all of which have well preserved shells, indistinct spiral cords to strong sigmoidal axial ribs crossed allowing microstructural details to be studied. by weaker spiral cords. Whorls with broad, smooth sutural Two Provanna specimens were collected from the Reki- ramp and distinct rounded shoulder, tabulated at axial ribs, fune whale-fall site in eastern Hokkaido (locality 2 in Fig. where present. 1; Amano et al. 2007). This site is from the Middle Miocene Description.—Medium-sized shell, up to 9.8 mm high, ovate Nupinai Formation (ca.13–12 Ma). The specimens are pre- fusiform; teleoconch at least three whorls; protoconch not pre- served as external moulds only, from which silicone rubber served. Sutures weakly impressed. Body whorl with broad, casts were made for examination. smooth sutural ramp sloping gently to rounded shoulder. A total of 124 Provanna specimens were collected from a From shoulder abapically body whorl sides gently curved. hydrocarbon seep deposit at the Kita-Kuroiwa Quarry in Ka- Ornament variable; some specimens, like holotype (Fig. 2A) kizaki-ku, Joetsu City, Niigata Prefecture, central Honshu, and paratype JUE 15902-2 (Fig. 2C), having whorls with Japan (locality 3 in Fig. 1; Amano et al. 2010). The Kuroiwa sigmoidal growth lines and many indistinct spiral cords best seep site is from the uppermost Middle Miocene (11.64±0.65 developed on the body whorl near the shoulder and at whorl Ma) Ogaya Formation (Amano et al. 2010). Twenty-eight of base; other specimens, like paratype JUE 15902-3 (Fig. 2D) the specimens are relatively well preserved and were exam- having nearly smooth whorls with sigmoidal growth lines ined in detail. The shells of these specimens are recrystal- only. Strong sigmoidal axial ribs on apical whorls of some lized, but relict microstructural details are visible in a few. specimens, such as paratype, JUE 15902-4, up to eighteen in To compare shell microstructural details of our fossil Pro- number on penultimate whorl (Fig. 2E). In some specimens, vanna species with modern species we studied the shell of distinct axial ribs also on body whorl, as in paratype, JUE AMANO AND LITTLE—MIOCENE PROVANNA FROM JAPAN 165 Table 1. Distribution, ecology, and shell microstructure of the fossil and modern species of Provanna. 1) after Kiel (2010). The fossil occurrence bathymetry estimations are in brackets because they are estimations. 2) The type material for these species was trawled and therefore the original habitats are unknown. The habitats given are from subsequent discoveries. 3) From inner layer (left) to outer layer (right); ccl, complex crossed lamellar structure; cl, crossed lamellar structure; homo, homogeneous structure; spl, simple prismatic structure. Height Shell Species name Age Geographic distribution Depth (m) Habitat (mm) microstructure3) Provanna tappuensis Cenomanian 3.9 Northwestern Hokkaido, Japan – seep Kaim, Jenkins, and Warén, 2008 Coniacian, seep and Provanna nakagawaensis 5.4 Northwestern Hokkaido – Kaim, Jenkins, and Hikida, 2009 Campanian wood-fall Eocene to seep and spl, ccl, homo Provanna antiqua 5.8 Washington State, USA (400–800)1) Squires, 1995 Oligocene wood-fall (Kiel 2006) Saether, spl, cl (Saether Provanna marshalii
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